Eur. J. Entorno!. 99: 315-351, 2002 ISSN 1210-5759

Generic revision and phylogenetic analysis of the Metriorrhynchinae (Coleóptera: Lycidae)

Ladislav BOCAK

Faculty of Science, Palacký University, tř. Svobody 26, 771 46 Olomouc, Czech Republic; e-mail: [email protected]

Key words.Coleóptera, Metriorrhynchmae, generic revision, phylogeny, taxonomy, key, World

Abstract. The subfamily Metriorrhynchinae is the most species-rich clade of Lycidae (Coleoptera). A recent proposal suggests that the Erotinae is a sister group of the Metriorrhynchinae. Within the Metriorrhynchinae, evidence is presented for the monophyly of the Conderini and Metriorrhynchini and their sister group position. The Trichalina, Hemiconderina and Metriorrhynchina form the tribe Metriorrhynchini. The relationships between the basal lineages of this group are poorly understood. Several clades are distin­ guished within the Metriorrhynchina, but there is only weak evidence supporting a relationships between them. The distribution of individual clades is discussed. Carathrix Kleine, 1926 (=Pseudodontocerus Pic, 1921), Dilolycus Kleine, 1926 (= Metriorrhynchus Gemminger et Harold, 1869),Flabelloporrostoma Pic, 1923 (= Metriorrhynchus Gemminger et Harold, 1869),Rossioptera Kasant- sev, 1988 (=Xylobanellus Kleine, 1930), Samanga Pic, 1921 (= Broxylus C.O. Waterhouse, 1879),Strophicus C. O. Waterhouse, 1879 (=Enylus C.O. Waterhouse, 1879), andTapromenoeus Bocak et Bocakova, 1989 (=Prometanoeus Kleine, 1925) are proposed as junior synonyms. Pseudosynchonnus Pic, 1922 is transferred to the Erotinae (Taphini) and Pseudosynchonnus Pic, 1922, Prota- phes Kleine, 1926, andParapyropterus Kleine, 1926 are proposed to be junior subjective synonyms ofLycoprogenthes Pic, 1915. Redescriptions of Metriorrhynchinae genera and a key to genera are provided.

INTRODUCTION first to introduce the genera of Metriorrhynchinae to lycid classification: Cladophorus Guerin-Meneville, 1830, The Metriorrhynchinae is the largest subfamily of Lyci­ Metriorhynchus Guerin-Meneville, 1838, andPorrostoma dae, both in the number of genera and number of species. Castelnau, 1838. Various authors disagreed with the Kleine (1933) listed in the last World Catalogue about dating of their descriptions, and the validity of these taxa 1200 species in 41 genera now classified in this was clarified much later by Bocak (1998c). Numerous subfamily. That is, in terms species this subfamily make authors added further species to various genera (e. g. up over 40 percent of the Lycidae. Most Metrior­ Erichson, Blanchard, Thomson, Walker, Motschulsky, rhynchinae are from Southeast Asia, the Indonesian Redtenbacher, Kiesenwetter, and Kirsch). C.O. Water­ Archipelago, New Guinea and Northern Australia. Fur­ house (1877, 1879) alone proposed thirteen new genera ther species occur in Africa, Madagascar, temperate Asia, which still form the backbone of Metriorrhynchinae clas­ Ceylon, and India. Only one species occurs in Eastern sification. He described also numerous new species from Europe and in Siberia. Within Lycidae, Metrior­ different parts of the Metriorrhynchinae range of distribu­ rhynchinae are defined by their circular phallobase (Figs tion. Further species were proposed in the last decades of 91-132), flat, conspicuous, unpaired vaginal gland in the XIXth century (Blackburn, Bourgeois, Fairmaire, most representatives (Figs 143-160), and a pronotum Gorham, Harold, Kirsch, Kiesenwetter, Lea, W. M. Leay, with lanceolate median areola, usually with lateral and and Schaufuss). Only two authors, R. Kleine and M. Pic, frontolateral pronotal carinae (Figs 57-86). Metrior­ contributed substantially to the knowledge of Lycidae rhynchinae appear to be close to the Erotinae as they have before World War II. They described hundreds of new very similar pronotal carinae in some forms, structurally species and several new genera. Kleine and Pic defined similar elytral costae, male and female genitalia similar in genera mostly on the basis of the presence or absence of some features, and generally similar larvae (Bocak and secondary costae, flabellate antennae or the number of Matsuda, in press). pronotal carinae. Congruence of these criteria with other Fabricius (1775) described the first species that is now characters needs to be established. Unlike Pic, Kleine dis­ placed in Metriorrhynchinae (Pyrochroa serraticornis sected genitalia, but preferred other characters for classi­ now classified in Trichalus) and later proposed several fication. Consequently, Kleine (1926a) classified additional species (Fabricius, 1801). Numerous entomolo­ Metriorrhynchus parallelus in Cladophorus on the basis gists working on the material collected by scientific expe­ of its flabellate antennae, although it is the type species of ditions in the first half of the XIXth century (Dalman, Metriorrhynchus, which he recognised. Fahraeus, Macleay, Hope, Klug, and Boisduval) fol­ Only a few species were described after World War II. lowed Fabricius. They described few species and classi­ Nakane (1968, 1969a, b, 1971, 1980) studied the fauna of fied them in then genera Lycus F., 1787, Calopteron Japan and Taiwan, Kasantsev (1988) described the genus Castelnau, 1838, orDictyoptera Latreille, 1829. Guerin- Rossioptera (= Xylobanellus Kleine) from Russia, but he Meneville (1830, 1838) and Castelnau (1838), were thedid not classify it in the Metriorrhynchinae. Bocak &

315 Bocakova (1987b, 1989, 1990b, 1991, 1999), Bocakobserved in early morning and shortly before sunset. (1998a, b, d, e, 1999a, b, 2000a, b, c, d, 2001a, b), andSome species are attracted to light but the proportion of Bocak & Matsuda (1998) published several geographi­ Lycidae collected at lights is generally low. Unlike Lyci­ cally limited revisions and descriptions of new species nae, the Metriorrhynchinae are rarely attracted to flowers. from the Oriental Region and New Guinea. Bocak Leptotrichalus are the only Metriorrhynchinae regularly (1998c) solved the long lasting uncertainty about the collected on flowers in South East Asia. Flowers in Aus­ status of the genera described by Guérin-Méneville. tralia attract at least some species of Porrostoma and Tri- Calder (1998) compiled a catalogue of Australian Lycidae chalus. Larvae were collected in decaying wood. Lycidae and proposed 85 new combinations within the Porros- generally seek moist conditions and inhabit rotten trunks toma. These new combinations were not based on the or branches in shaded places, often laying on the ground study of type specimens, and therefore the generic classi­ and in close contact with moist soil. Several larvae of fication of Australian Lycidae remains obscure. Metriorrhynchinae have been described. Burakowski At present, the entire classification of the Metrior- (1988) described the larva and pupa Xylobanellus of rhynchinae at the species and generic levels is chaotic. A erythropterus; Hayashi (1954), Fukuda & Hayashi large part of this confusion results from the inadequate (1981), and Hayashi (1986) described several Japanese nature of Pic’s descriptions and unclear reasons for his species of Cautires. Potozkaja (1981) described the larva proposals. He thought that as his descriptions were long of Xylobanus angusticollis (Motschulsky). Other larvae they would be useful (W. Wittmer, pers. comm.). As of Metriorrhynchinae were reported recently by Bocak & Kleine usually ignored Pic’s descriptions, revisions of Matsuda (in press). Metriorrhynchinae are often involved most genera are urgently needed. in Mullerian mimicry complexes, Lawrence & Britton The subfamily Metriorrhynchinae was proposed by (1991) mentioned those from Australia. Moore & Brown Kleine (1926a), along with the subfamily Dilolycinae (1981) studied the antifeedants in the bodyof Porrostoma (=Haplothoracinae, nomen nudum). Later he described rhippidium (W. M. Leay, 1827). the subfamilies Trichalinae and Cladophorinae (Kleine, MATERIAL AND METHODS 1928). Kleine (1933) designated all these taxa as tribes of Lycinae. Bocak & Bocakova (1990a) proposed the tribe Taxa Conderini and subtribe Hemiconderina, synonymised Altogether 45 genera were classified in the Metriorrhynchinae Cladophorinae and Dilolycinae with Metriorrhynchinae and Bocakova (in press) transferred Broxylus to the Metrior­ and downgraded Trichalinae to the subtribal level. Metri­ rhynchinae: for a list of genera see Appendix 1. The phyloge­ orrhynchinae was made up of two tribes, Conderini and netic analysis deals with 34 genera of Metriorrhynchinae. One Metriorrhynchini, the later being divided to the subtribes of Metriorrhynchinae genera was found to be a member of the Erotinae (Pseudosynchonnus). Four genera were synonymised Trichalina, Metriorrhynchina and Hemiconderina. Bocak prior the analysis, because they did not differ from genera (1998b) revised the generic classification of the Tri- included in the analysis: Dilolycus (= Metriorrhynchus), Flabel- chalina. loporrostoma (= Metriorrhynchus), Samanga (= Broxylus), and The species richness and high endemism in the transi­ Strophicus (= Synchonnus). One genus was not found in Pic’s tional Austro-Oriental zone, as well as the restriction of collection and its identity is unknown (Falsolucidota). The Lycidae to moist forests and usually to mountainous habi­ genera Mimoxylobanus, Oriomum, Cladophorinus, Xylobano- tats, suggest that Metriorrhynchinae could be a valuable morphus, Xylobanomimus, and Malacolycus are available only model group for evolutionary and zoogeographical stud­ as single specimens of one sex, and therefore they were ies. This characteristic is combined with the fact that excluded from the analysis. Their redescriptions and discussion Lycidae are models in numerous mimicry complexes. of their possible systematic position are given in the taxonomic part. The proportion of unknown character states is very high Despite all this, the poor knowledge of Metriorrhynchinae for these genera and this uncertainty collapsed the basal part of has prevented them from being used for more general the tree of Metriorrhynchini to the unresolved bush. Wiens studies. Research on this group has suffered from the (1998) discussed the consequence of a high proportion of absence of comprehensive definitions of genera as well as missing data. All the genera excluded from the analysis as well the doubtful phylogenetic affinities between genera and as genus of unknown identity are monotypic, therefore their tribes. This paper should establish the basis for further exclusion did not restrict substantially the scope of the analysis. study of this extremely diverse group. All type species of Metriorrhynchinae genera were studied except Falsolucidota Pic, 1921b, which should be deposited in Natural history the National Museum of Natural History in Paris, but was not Metriorrhynchinae are one of the commonest groups of found in Pic’s collection. Lycidae in tropical Asia and Australia. The adults live Bocak & Bocakova (1990a) reclassified the Metriorrhynchi­ mostly on the leaves of shrubs and herbs in dense forest nae. Evidence for monophyly was sought in the morphology of canopy. Although most specimens in museum collections the ingroup and trees were rooted using the outgroup method (Watrous & Wheeler, 1981). A sister-group for the Metrior­ were collected individually or by sweeping of the lower rhynchinae was not suggested by previous authors and the pos­ strata of tropical forests, Metriorrhynchinae are active sibility of a close relationship between Erotinae and also in the upper strata as shown by the material obtained Metriorrhynchinae based on larval characters was briefly dis­ by insecticidal fumigation in Sulawesi (Bocak, 2000c). cussed by Bocak & Matsuda (in press). In addition, Erotinae and They are generally slow moving and fly only occasionally Metriorrhynchinae have the same arrangement of elytral costae, and reluctantly. The highest dispersal activity was carinae on the pronotum and slender parameres (present only in

316 Conderis Conderis 1 Xylobanellus 2 Xylobanellus Synchonnus Enylus Achras Diatrichalus Hemiconderis Trichalus Wakarumbia Microtrichalus Enylus Flabellotrichalus Diatrichalus Schizotrichalus Flabellotrichalus Eniclases Schizotrichalus Synchonnus Eniclases Achras Trichalus Flemiconderis Microtrichalus Wakarumbia Metriorrhynchus Bulenides Leptotrichalus Caenioxylobanus Lobatang Cautires Metanoeus Xylobanus Stadenus Broxylus Metriorroides Prometanoeus Porrostoma Tapromenoeus Kassemia Stadenus Carathrix Metriorroides Pseudodontocerus Porrostoma Cladophorus Metriorrhynchus Procautires Leptotrichalus Cautiromimus Lobatang Ditua Metanoeus Bulenides Cladophorus Caenioxylobanus Procautires Cautires Cautiromimus Xylobanus Ditua Broxylus Kassemia 25 Prometanoeus Carathrix HL Tapromenoeus Pseudodontocerus

Figs 1-2: Cladograms. 1 - the strict consensus tree derived from three trees provided by an analysis of the matrix in Table 2 (mh*, bb*); 2 - the strict consensus tree based on two trees obtained after application of successive weighting. The numbers above branches correspond to node numbers in Tab. 3.

Conderini within the Metriorrhynchinae). The phylogeny of Characters Lycidae has not yet been analysed, and therefore multiple out­ I identified 72 characters, 67 potentially informative for the groups were chosen and the impact of switching to different out­ relationships within an ingroup and five of them constant within groups tested. The following taxa were used as outgroups: an ingroup and used for its definition. Polymorphic characters in Dictyoptera (represented by D. aurora (Herbst)) andPyropterus terminal taxa were coded as “-” in the matrix, and all characters (P. nigroruber (Degeer)) from Erotinae as presumably the are described in Table 1. Autapomorphic characters of indi­ closest sister-group, Lyponia (L. (Ponyalis) laticornis vidual genera were not included and they are discussed in the Fairmaire), which is considered to be the most basal member of taxonomic part. When possible, several species of each genus Platerodinae (Bocakova, 2001), and Calochromus were scored for each character. The genera included in the (Calochromus sp. from Yunnan) from the Calochrominae. Dic- analysis were always available as type(s) or identified speci­ tyoptera and Pyropterus represent genera that have and lack mens of type species of both sexes. In several cases, data from secondary elytral costae. This character was thought to be disarticulated closely related species were compared with the important for the generic classification of Lycidae. The selected type species. Because of the danger of inappropriate sampling, outgroups represent all potential relatives of Metriorrhynchinae. the examination of several representatives of each genus was The remaining subfamilies of Lycidae differ substantially in preferred. This was easy in case for recently revised genera, but adult and larval characters (Lycinae: Lycini and Macrolycini) difficult in the case of the very large genera used by M. Pic and in addition some of them have neotenous females ( Duliti- (Metriorrhynchus, Cladophorus, Cautires), for genera typologi­ cola, Platerodrilus, presumably Ateliinae, Leptolycinae: Dexo- cally consisting of very diverse groups of unrelated species rini). (Procautires) or genera formally monotypic but based on numerous undescribed species. I preferred to score the group of species closely related to the type-species. An alpha-taxonomic study of such genera was beyond the scope of this paper.

317 Figs 3-6: Distribution. 3 - distribution of subfamily and tribes; 4 - eastern limit of range of Asian continental elements; 5 - distri­ bution of Hemiconderina and Trichalina; 6 - western limit of range of Australian elements.

The present study is based exclusively on adults. Both male screen or illustrations were derived from photographs taken with and female genitalia of all available type specimens were dis­ an Olympus DP-10 or Camedia 3000 digital camera attached to sected and the membranous parts stained lightly with chlorazol an Olympus SZX-12 stereoscopic microscope. black. Important characters were drawn using an ocular grid-

318 Figs 7-28: 7-9. General appearance. 7 -Enylus amplus; 8 - Wakarumbia grandis; 9 - Lobatang sordescens; 10-11. Head. 10 - Porrostoma rhippidium; 11 - Diatrichalus sp.; 12 - pronotum, Metriorrhynchus fessus; 13 - labrum,Pseudodontocerus pulcher; 14 - labium, Metriorrhynchus parallelus; 15-17. Maxilla. 15 - Broxylus fenestratus; 16 - Prometanoeus ochraceus; 17 - Metanoeus sp.; 18 - maxillary palpus,Pseudodontocerus pulcher; 19-21. Labrum and hypopharynx. 19 - Metriorrhynchus parallelus; 20 - Conderis signicollis; 21 - Metanoeus sp.; 22-23. Scutellum. 22 - Schizotrichalus nigrescens; 23 - Metriorrhynchus mirabilis; 24-28. Mandible. 24 -Metriorrhynchus parallelus; 25 - Conderis signicollis; 26 - Broxylus fenestratus; 27 - Pseudodontocerus sp.; 28 -Cladophorus sp. Scale 0.5 mm (Figs 1-6, 17), 0.2 mm (Figs 7, 12), 0.1 mm (Figs 8-11, 13-16, 18-22).

319 Table 1. List of characters and character states used in the cladistic analysis of Metriorrhynchinae 1 Rostrum 0, absent (Fig. 11); 1, present (Fig. 10) 2 Clypeo-genal suture 0, absent or unclear; 1, short; 2, long 3 Gula 0, longitudinal or rectangular; 1, transverse (Fig. 11) 4 Frontal margin of clypeus 0, convex; 1, concave; 2, straight; 3, bisinuate; 4, shallowly emargi- nate 5 Apexoflabrum 0, simply rounded; 1, emarginate (Figs 19-21) 6 Labral arms 0, long, slender; 1, with stout base; 2, fused with lateral margin of labrum 7 Mandibles 0, long, stout, considerably curved (Fig. 25, 27, 28); 1, small, slender (Fig. 24); 2, small, slender with apex curved at a right angle 8 Molar 0, present, or at least mandibular base robust (Figs 25-28); 1, absent (Fig. 24) 9 Ventral mandibular condyle 0, ventrally directed; 1, caudally directed, robust; 2, caudally directed, weak 10 Apical segment of maxillary palpus 0, robust (Fig. 15); 1, slender (Fig. 16-17) 11 Tubercles on margin of terminal maxillary palpomere 0, absent (Fig. 15-17); 1, present (Fig. 18) 12 Tubercles on margin of terminal labial palpomere 0, absent (Fig. 14); 1, present 13 Antennae 0, rounded in cross-section; 1, considerably compressed 14 Maleantennomeres3-10lamellae 0, with wide lamellae, serrate, filiform (Figs 29-39); 1, with slender lamellae 15 Maleantennomeres3-10 0, filiform or serrate; 1, flabellate (Fig. 38) 16 Length of antennomere 2 0, comparable with antennomere 3; 1, considerably shorter (Fig. 29-51) 17 Femaleantennae 0, serrate or filiform (as in Fig. 30); 1, flabellate (Figs 46-47) 18 Frontolateral pronotal carinae 0, absent (Figs 57-65); 1, present or at least vestigial (Figs 69-83) 19 Lateral pronotal carinae 0, present (Figs 57-59, 69-83); 1, absent or vestigial (Figs 61-68) 20 Frontal margin of pronotum 0, simple Figs 57-66, 69-86); 1, strengthened, formed by two carinae (Figs 67-68) 21 Medianareola 0, constricted at base; 1,not constricted (Figs 87-86) 22 Median areola/frontal margin 0, directly attached (Fig. 61); 1, connected by costa (Figs 57-59, 62-86) 23 Shape of median areola 0, in middle, usually large (Figs 57-61); 1, small, short, attached to pronotal base (Figs 68-73, 77-86); 2, slender,Trichalus type (Figs 62-65) 24 Base of pronotum 0, with symmetrical bulges; 1,without symmetrical bulges 25 Number of pronotal areoles 0, less than seven (Figs 57-68); 1, seven (Figs 71-83) 26 Shape of pronotum 0, with apparent frontal angles and elevated lateral margins (Figs 57-73, 75-80); 1, rounded frontal angles, compact lateral margins (Figs 74,81) 27 Pronotal carinae 0, robust, obtuse; 1, slender, straight (Figs 59-60) 28 Lateral margins of pronotum 0, without tubercles (Figs 57-59); 1, with tubercles (Figs 61-66); 2, slightly widened before base (Figs 76, 78) 29 Secondary elytral costae 0, fully developed (Figs 52-55); 1, present only close to humeri; 2; completely absent (Fig. 87); 3; absent only in middle part 30 Primary elytral costa 1 0, fully developed (Fig. 8); 1; considerably shortened (Fig. 9) 31 Primaryelytralcosta3 0, full; 1, weaker at apex; 2; considerably shortened; 3, absent 32 Strength of costae 0, equal; 1, costae 2 and 4 much stronger, or at least costa 2 stronger 33 Elytral cells 0, transverse or rectangular (Fig. 55); 1, rounded, small (Fig. 54) 34 Process of mesothoracic spiracular plate 0, absent; 1, short; 2, long; 3, without process, spiraculum in form of large tube

35 Metascutellar hind margin 0, emarginate with keel; 1, straight without keel; 2, straight with keel 36 Mesoscutellar lateral process 0, absent or short; 1, long 37 Lenght of alacrista 0, as long as scutoscutellar suture or longer; 1, distinctly shorter

320 Table 1 (continued). 38 Width of mesostemal process 0, wide; 1,narrow 39 Tibial spurs on hind legs 0, present; 1, absent; 2, very short 40 Apical part of phallus 0, rounded in cross section or dorsoventrally compressed; 1, consid­ erably laterally compressed 41 Phallus short, wide, robust 0, no; 1, yes (Figs 122, 125, 127) 42 Phallus lanceolate, widest at apex 0, no; 1, yes (Figs 92, 107, 108, 109) 43 Phallus slender, narrows at apex, membranous ventrally 0, no; 1, yes (Figs 103, 104) 44 Phallus slender, tubular 0,no; 1,yes(Figs 117-120) 45 Internal sac 0, membranous, without sclerotized structures (Fig. 91); 1, with two apical paired spines (Figs 107-110); 2, with two basal paired spines (Figs 125, 127); 3, linearly arranged spines; 4, spines fused (Fig. 94); 5, with stick-like structure (Fig. 121); 6, phallus with unique struc­ ture apically (Figs 100-101) 46 Internal sac with sclerotised plates or ducts 0, no; 1, yes (Figs 121-123, 125-126) 47 Phallus very slender, with weakly bulbous base, apex 0,no; l,yes (Fig. 94) often rotated 48 Process at bases ofparamerae 0, absent; 1,present 49 Phallus with dorsal keel 0, no; 1, yes (Figs 105, 106) 50 Base of phallus 0, straight; 1, hook-like (Fig, 91) 51 Basal tubercle of phallobase 0, absent; 1,present 52 Internal sac 0, completely membranous, colourless, retracted in phallus (Fig. 91); 1, exposed, at least partly sclerotized or coloured 53 Primary gonoporus 0, membranous; 1, sclerotized (Figs 122, 125, 126) 54 Parameres 0, present; 1,absent (Figs 92-132) 55 Shape of basal partof phallus 0, fused with basal part of parameres; 1, simply narrowed (Fig. 112); 2, obtuse orwidened (Fig. 121, 127] 56 Symmetry of basal partof phallus 0, symmetrical (Fig. 112); 1, asymmetrical, type Metriorrhynchus (Fig. 114-116); 2,asymmetricaltype Trychalus (Fig. 104) 57 Phallobase 0, elongate; 1, circular 58 Basal margins of coxites 0, strengthened; 1, simple; 2, plate-like 59 Phallobasal membrane 0, soft, colourless; 1, pigmented, sometimes sclerotised (Figs 114-116) 60 Valvifers fused with coxites 0, no; 1, yes (Fig. 156, 159) 61 Valvifers distinctly widened medaslly 0, yes; 1, no (Fig. 133) 62 Valvifers 0, much longer than coxites; 1, moderately long (Figs 151-153); 2, shortened, widened at apex (Fig. 136); 3, vestigial (Fig. 150) 63 Lateral glands with narrow ducts 0, no (Fig. 142); 1, yes (Figs 147-160) 64 Unpaired gland 0, absent or minute, filiform (Fig. 142); 1, flat, distinct (Fig. 147) 65 Lateral glands 0, inserted apically or laterally (Fig. 147); 1; inserted dorsally 66 Spermatheca 0, bulbous at least apically (Figs 142-144); 1, very slender 67 Paraproctus 0, with lateral sclerites fused to valvifers; 1, with free lateral sclerites; 2, simple, without lateral sclerites 68 Basal part of male sternum S8 0, very narrow basally; 1, quite wide, simply rounded; 2, wider, short; 3, widened, constricted before apex 69 Sternum S7 0, emarginate; 1, straight 70 Window in male sternum S8 0, present; 1, absent 71 Spiculumgastrale 0, present (Fig. 162); 1, absent 72 Apex of female sternum S7 0, simple; 1, shallowly emarginate; 2, deeply emarginate

Phylogenetic analysis analyses. The results obtained by this procedure are uncertain to A total of 72 characters, as given in Table 2 were run unor- be of minimal length t a the implicit enumeration was not used dered and unweighted. The minimum length trees were searched because the matrix was too extensive and the computer equip- using computer program Hennig86 ver. 1.5 (Farris, 1988) using ment unable to deliver the results in a reas°nable time. Subse­ the Wagner parsimony analysis. The procedure of “mh*” (mul- quently using the xstep command each character was tiple hennig) and “bb*” (branch breaking) were used in all weightedaccording to its fit to the tree in the preceding ran. The

321 Figs 29-56: 29-51. Basal antennomeres. 29 - Conderis signicollis, male; 30 - Achras limbatus, male; 31 - Hemiconderis gracilis, male, ventral view; 32 - Diatrichalus salomonensis, male; 33 - Diatrichalus assimilis, male; 34 - Microtrichalus griseus, male; 35 - Trichalus flavopictus, male; 36 - Metriorrhynchus mirabilis, female; 37 - Prometanoeus ochraceus, male; 38 - Cautires ratus, male; 39 - Caenioxylobanus ater, female; 40 - Metanoeus pallidus, male; 41 - Metriorrhynchus inaequalis, male; 42 - Porrostoma lateralis, male; 43 - Metriorrhynchoides helleri, male; 44 - Cladophorus elegantulus, male; 45 - Pseudodontocerus sp., male; 46 - Pseudodontocerus pulcher, female; 47 - Pseudodontocerus sp., female; 48 -Malacolycus paululus, male; 49 - Porrostoma rhipid- ium, male; 50 - Mimoxylobanus reticulatus , male; 51 - Cladophorus sp., male; 52-56. Elytral reticulation. 52 - Porrostoma sp.; 53 - Pseudodontocerus pulcher; 54 - Pseudodontocerus sp.; 55 - Metriorrhynchoides sp.; 56 - Malacolycus paululus. Scales 0.5 mm.

322 Table 2. Data matrix 000000000111111111122222222223333333333444444444455555555556666666666777 123456789012345678901234567890123456789012345678901234567890123456789012

Dictyoptera 000000001000000000000000000000000000000000000000000000000000000000000001 Calochromus 00020001200000000010--- 00000-- --0000000000000000000000000000100000000102 L yp o n ia 000110001000-0100010---0000000100010000000000000000-01100200010000010002 P y ro p teru s 000100001000000000000000000010000000000000000000000000000000000000000100 C o nderis 011010001000-00100001101000000000100000100000001011000001000011000001002 Xylobanellus 011110001000-00100001101000020000100000100000001011000001000011000001002 Diatrichalus 02111001001110-1001011210001-1000101101000006000000101111100111101210101 E n y lu s 001110011100100100101-010001002101111110000020-0000101121100111101220111 Flabellotrich. 020310212100101100101121000101000111101000100000000101121100111110220101 Schizotrichalus 0203102121001001011011110001010002-1101000100000100101121100111110220101 E n icla ses 00031021210010-101101--1000101000111101000100000100101121100111110220101 T rich a lu s 020310212100100100101121000101000201101000102000000101121100111110230100 Microtrichalus 000310212100100100101121000101000201101000102000000101121100111110220100 S yn c h o n n u s 021110010001100100001001000000110101101001001000000101201100111100120110 A c h ra s 011110200001100100001101001000110201101001001000000101201100111100120111 Hemiconderis 021110200001100100001101001000210201101000003010000101201100111100220111 W akarum bia 021110200001100100001101001010110101111000004010000101201100111100230111 B u len id e s 021110000100101101111111000000000211102001001000000101101100111100120100 Caenioxylobanus 021110000001101100111111000020000211101001001000000101101100111100110100 C a u tires 0111000001001011010- 111110000000020110200100100000010110110011110012010- X y lo b a n u s 01110000100010- 101-011111000200002-110200100100000010120110011110011010- B ro xy lu s 011110000000100101-01111100020000211101001001000000101101100111100210100 Prometanoeus 011100010100100101001111100000300001101001001000000101101100111100120100 Tapromenoeus 011100010100100101001111100020300011101001001000000101101100111100120100 M eta n o eu s 021110010100111101001111100-001002111000-0002000000101201110131100220111 S ta d e n u s 021400011101101101-01111-002001-0411100000010000000101101101111100220110 Metriorrhynchoid. 100412112100101101001111100200000420000000010000000101101101111100120111 P o rro sto m a 100412112100101101001111100200100420100000010000000101-0110111110022011- Metriorrhynchus 02101010010010-101001111110000000210000000010000000101111100111100130110 Leptotrichalus 02020210210010010010112100000100021110100001000000010111111012110---0111 L o b a ta n g 020202102100100100101121000001000211111000010000000101111110121100130111 Cladophorus 0214100000001-1101001111100-00000211110000005100000111211110111100130110 Procautires 021110000000101101001111100-30000211111010000100000111101110111100130100 Cautiromimus 011001000001101101001111110000000211111010000100000111-0110011110013011- K a sse m ia 011111001001101101001111100021100111101010002100000111201100111100221100 D itu a 011001000100100101001111100000001211110010002100000111201100111100130100 C a ra th rix 021001000100111111001111110000001-01101010002100000111201110111100120111 Pseudodontocerus 021011001011111111001111110000001101101010002100000111101110111100120111 weights were reassigned until the topology of the resultant trees steps, a Cl = 0.39 and a RI = 0.69. The strict consensus did not change. The program Tree Gardener ver. 2.2 (Ramos, tree derived from them was identical with one of the 1997) was used for matrix edition and control of Hennig86 pro­ original trees and included a polytomy in Trichalina (Fig. gram. The Clados ver. 1.9 program (Nixon, 1998) was used for 1), and therefore the length, CI and RI of these trees did mapping of characters on a tree and a program default setting not differ. When only one of the original outgroup taxa “homO” was applied. This setting indicates as homoplasious only those changes that are designated as of independent origin. was designated as an outgroup, the topology of the tree did not change. The mapping of characters is given in Depositories table 3, and the number of clades in Fig. 1. Material was examined from the following institutions and Further, successive weighting was applied to the matrix collections: BMNH, Natural History Museum, London; BPBM, after mh* and bb* (again with Calochromus, Lyponia, State Museum of Natural and Cultural History, Honolulu, Dictyoptera and Pyropterus as outgroups). Two trees Hawaii; DEIE, Deutsches Entomologisches Institut, Eberswalde; HNHM, Hungarian Natural History Museum, with length 651, a CI = 69 and a RI = 86 were obtained. Budapest; MHNG, Muséum national d’histoire naturelle, These trees differed in the position of Xylobanus, either Geneva; MHNP, Muséum national d’Histoire naturelle, Paris; being a sister-group of Cautires or Broxylus. The con­ MNHU, Museum für Naturkunde, Humboldt Universität, sensus tree was slightly longer (length 653) and had iden­ Berlin; MHNB, Museum d’Histoire naturelle, Brussels; MSNG, tical indexes. The strict consensus tree differs in that it Museo civico di storia naturale, “Giacomo Doria”, Genova; has a polytomy in the Cautires clade. NHMB, Naturhistorisches Museum, Basel; NHMW, Naturhis­ torisches Museum, Vienna; RMNH, Natural History Museum, Discussion Leiden; SMNS, Staatliches Museum für Naturkunde, Stuttgart; The present analysis is the first attempt to investigate SMTD, Staatliches Museum für Tierkunde, Dresden; USNM, the phylogeny of the Metriorrhynchinae using a cladistic National Museum of Natural History, Washington, D.C.; approach. The only classification of Lycidae (Bocak & ZMAN, Zoological Museum, Amsterdam; ZMPA, Instytut Zoo­ Bocakova, 1990a) is not based on phylogenetic principles logii PAN, Warsaw; LMBC, author’s collection. but on the need to diagnose supergeneric taxa for a com­ PHYLOGENETIC ANALYSIS parative morphology of the major groups of Lycidae, and their identification. Nevertheless, the monophyly of the Results Metrirrhynchinae as understood by Bocak & Bocakova When Calochromus, Lyponia, Dictyoptera, and Pyrop- (1990a), as well as their constitutive tribes and subtribes terus were used to root cladograms, the analysis (mh*, is supported by the present phylogenetic analysis. The bb*) yielded three parsimonious trees, each with 261

323 Figs 57-90: 57-86. Pronotum. 57 - Xylobanellus erythropterus; 58 - Conderis signicollis; 59 - Synchonnus clientulus; 60 - Hemi- conderis explicatus; 61 - Enylus amplus; 62 - Diatrichalus salomonensis; 63 - Microtrichalus acutangulus; 64 - Microtrichalus patricius; 65 - Flabellotrichalus bilineatus; 66 - Eniclases nigriceps; 67 - Caenioxylobanus ater; 68 -Bulenides inapicalis; 69 - Xylobanus benignus; 70 - Broxylus fenestratus; 71 - Cautires ratus; 72 - Prometaneus ochraceus; 73 - Metanoeus dispar; 74 - Metriorrhynchus thoracicus; 75 - Leptotrichalus accuratus; 76 - Stadenus puncticollis; 77 - Porrostoma lateralis; 78 -Metrior- rhynchoides helleri; 79 - Cladophorus praecipuus; 80 -Ditua deplanata; 81 -Cautiromimus reticulatus; 82 -Procautires toxopei; 83 -Pseododontocerus pulcher; 84 - Malacolycus paululus; 85 - Dilolycus lamellatus; 86 -Mimoxylobanus reticulatus; 87-90. Elytral reticulation. 87 - Xylobanellus erythropterus; 88 -Caenioxylobanus ater; 89 -Cautires sinensis; 90 - Broxylus fenestratus. Scales 0.5 mm.

324 Table 3. Synapomorphies and homoplastic changes listed by nodes for the cladogram in Fig. 1. Nodes Synapomorphies Homoplastic changes 1 Metriorrhynchinae 3.1, 13.1, 16.1,21.1,24.1,57.1 34.1 2 Conderini 40.1,48.1,50.1,51.1 2.1, 69.1 3 Metriorrhynchini 2.2, 9.0,36.1,37.1,39.1,52.1 54.1,55.1,61.1,67.1 58.1, 63.1, 64.1, 68.2 70.1,71.1,72.1 4 Hemiconderina - 12.1,31.1,32.1,45.1,55.2 5 27.1 7.2 6 47.1 67.2 7 - 10.1, 35.1 8 Trichalina 28.1, 56.2 8.1, 19.1, 67.2 9 - 23.2, 30.1,71.0 10 4.3,43.1, 65.1 3.0, 7.2, 9.2 11 - 34.2,35.0,45.2, 72.0 12 49.1 18.1,23.1 13 Metriorrhynchina - 23.1, 34.2 14 - 4.0,7.1,44.1,56.1,68.3 15 62.2 3.0, 4.2, 5.0, 6.2, 9.2, 19.1, 23.2, 30.1,59.1 16 - 15.1, 18.1,25.1 17 - 8.1,31.1,39.0, 67.2 18 Porrostoma-group 28.2, 34.4, 60.1 4.4, 44.1 19 1.1,35.2 2.0,3.0,6.2,7.1,9.2,36.0 20 - 71.0, 72.0 21 - 42.1,45.1 22 Bulenides-group 20.1 19.1,25.0 23 Cautires-group - 2.1, 5.0 24 - 15 25 31.3 8.1, 34.0 26 - 10.0, 29.2, 68.1 27 Ditua-group 41.1,46.1,53.1 10.0, 55.2 28 - 6.1,34.1,45.2 29 17.1 4.0, 14.1, 26.1, 33.1, 35.0, 59.1, 71.1, 72.1 30 - 38.1, 68.3 31 - 2.1, 4.0, 5.0, 6.1 32 - 59.1 monophyly of the Cladophorini as defined by Kleine antennomere 2 as have several Leptolycinae. Both groups (1933) and elsewhere was rejected by both hypotheses differ substantially in the shape of their cranium, male (Figs 1, 2). genitalia and Ateliinae have pobably neotenous females The monophyly of the Metriorrhynchinae is supported (Bocak & Matsuda, in press). These groups are unrelated by several synapomorphic characters: transverse gula to the Metriorrhynchinae, therefore an independent origin (Fig. 11), considerably compressed antennae, antenno- of these character states in the Metriorrhynchinae is sug­ mere 2 much shorter than antennomere 3 (Figs 29-51), gested. unconstricted base of median areola (Figs 57-86), median The Conderini, according to the present phylogenetic areola connected to frontal margin of the pronotum by a hypothesis (Figs 1, 2), have a basal placement within the keel (Figs 57-60, 62-86), symmetrical bulges at base of Metriorrhynchinae. That the phallus has a laterally com­ pronotum absent, circular phallobase (Figs 91-132). The pressed apical part, and a hook-like base, and there are shape of the phallobase is very characteristic and unlike processes on the phallobase support their monophyly. All that of other Lycidae. In addition, the shape of the pro­ these characters are unique within the Lycidae. Addition­ notal carinae is very characteristic and only specimens of ally, the shape of the pronotal carinae is very character­ Enylus is these an indistinct frontal keel, but even in such istic in most species (Figs 57, 58), but the shape of the cases the general appearance of the pronotum is different. carinae is not the same in all specimens of some species The very short antennomere 2 is possibly a synapomor­ and therefore was not coded in the matrix. Conderini phic character state, although Ateliinae have a very short share several symplesiomorphic character states with the

325 Figs 91-111: Male genitalia. 91 - Conderis signicollis; 92- 93 -Synchonnus clientulus; 94 - Wakarumbia angustior; 95- 96 - Achras limbatus; 97 - Enylus segregatus; 98 - 99Enylus - amplus; 100 -101 - Diatrichalus assimilis; 102 - Diatrichalus sp.; 103 - Microtrichalus bicoloripes; 104 - Flabellotrichalus notatithorax; 105 - Schizotrichalus nigrescens; 106 - Eniclases apertus; 107 - Bulenides sp.; 108 -Cautires sinensis; 109 -110 -Xylobanus costifer; 111 - Broxylus fenestratus. Scales 0.5 mm.

Erotinae. The female genitalia have a very similar shape The monophyly of the clade forming the tribe Metrior- and the basal part of coxites is streghthened, male geni­ rhynchini is supported by the regularly exposed, sclero- talia have long parameres, and a completely membranous tized or at least pigmented internal sac (Figs 92-132), the internal sac. flat, apparent, unpaired vaginal gland (e.g. Figs

326 Figs 112-130: Male genitalia. 112 - Prometanoeus ochraceus; 113 - Metanoeus sp.; 114 - Metriorrhynchus thoracicus; 115 - Leptotrichalus adolescens; 116 - Lobatang dubitabilis; 117 -118 -Stadenus triareolatus; 119 - Porrostoma rufpennis; 120 - Met- riorrhynchoides helleri; 121 - Cladophorus sp.; 122-123 - Cautiromimus reticulatus; 124 - Cladophorinus cyanescens; 125 - Ditua deplanata; 126 - Procautires sp.; 127-128 -Pseudodontocerus sp.; 129 - Malacolycus paululus; 130 - Xylobanomorphus trans- formis. Scales 0.5 mm.

143-146), lateral vaginal glands with a narrow, usually Tab. 3, and the characters not mentioned here have under­ slightly sclerotized base (e. g. Figs 143-146), simple gone multiple reversions. The absence of parameres are basal margin to coxites (Figs 134-140), mesoscutellum hypothesised to be a homoplasy, because this character with long lateral processes, and short alacrista. The com­ state has evolved independently in Lyponia (Platerodinae, plete set of synapomorphic character states is given in char. 54) and is known also in the Platerodini, which were

327 not included as an outgroup in the analysis. Metrior- 18.1, after successive weighting (Fig. 2) the state 18.1 is rhynchini differ in this character from all similar Lycidae. not shared by the basal Metriorrhynchus+Lobatang+Lep- However, the loss of parameres has occurred independ­ totrichalus clade. Additionally, this character is a homo- ently in other groups. plasy as it is a similar arrangement of carinae in The tribe Metriorrhynchini is formed by three subtribes: Schizotrichalus and Eniclases (Fig. 66). The arrangement Trichalina, Hemiconderina and Metrirrhynchina. Their of carinae in Eniclases is quite different from that of most relationship is unresolved. When no successive weighting Metriorrhynchina - only two diverging keels are present was applied, the Trichalina had a basal position as a (Fig. 66). Schizotrichalus has similar longitudinal carinae sister-group of the remaining Metriorrhynchini, and the but its median frontal carina is forked apically and con­ Hemiconderina were the next basal clade. After succes­ nected with longitudinal carinae. Consequently, three sive weighting the relative position of these subtribes carinae are attached to the frontal pronotal margin. changed (Figs 1, 2). But in all analyses these clades Having no evidence to the contrary, both situations were remained stable. Unfortunately the larvae of these groups scored as an identical character state. Further evidence are unknown and I have not found a character to corrobo­ will be needed to support or refute the Metriorrhynchina. rate the hypotheses. Both hypotheses can be interpreted in As few larvae of the Metriorrhynchina have been the same way in the formal classification. described (Bocak & Matsuda, in press) they will not help Kleine (1928, 1933) defined the Trichalina as a group to solve this ambiguity. The larvae of most known Cau- of Metriorrhynchinae genera with a short elytral primary tires and Xylobanus are very similar and they share some costa 1. Bocak (1998b) showed that the shortened costa 1 characters with Metriorrhynchus. Porrostoma differs in evolved several times in the Metriorrhynchinae and there­ having characteristic processes on the hind margins of its fore restricted Trichalina to the terminal clade Trichalus + terga. Most of them share with Xylobanellus terga that are Microtrichalus + Flabellotrichalus + Schizotrichalus + divided in two parts by a membranous line. Eniclases and provisionally classified Enylus and Diatri- Only two constitutive clades give robust support to both chalus as incertae sedis in the Metriorrhynchini. The pre­ cladograms (Figs 1,2): the Ditua group formed by almost sent analysis placed Enylus and Diatrichalus basally in exclusively Papuan genera and the Porrostoma group this clade because of two synapomorphic character states: with a restricted Austro-Papuan distribution. The clade tubercles on the lateral margin of the pronotum (Figs Bulenides + Caenioxylobanus is either a basal group (Fig. 61-66) and the shape of the basal part of phallus (Figs 1) with the Cautires clade as the next clade, or they are a 104, 106). The tubercles may or may not be distinct in basal group within the Cautires clade (Fig. 2). individual specimens but are regularly present, and The position of Metanoeus is variable and it is attached together with the characteristically elevated lateral mar­ as a basal member either to the Porrostoma or Ditua gins give the uniform pronotal appearance of the Tri­ clade. In any case, Metanoeus, has an Oriental distribu­ chalina. Considering the topology of the trees produced tion, and is not related to the Afro-Oriental clade (Cau­ by the present analysis (Figs 1, 2), Enylus and Diatri­ tires, Bulenides, Prometanoeus, etc.). Some characters chalus must be classified in the Trichalina. present only in some members of these genera could not The Hemiconderina as proposed by Bocak & Bocakova be coded in the matrix: several Metriorrhynchus species (1990a) was monotypic. Later, Bocak (1999b) described share with Metanoeus very obtuse, indistinct pronotal Wakarumbia from Sulawesi put it in the Hemiconderina. carinae and both have shorter (Metriorrhynchus) to ves­ This was supported by the present analysis and addition­ tigial (Metanoeus) valvifers. Further evidence is needed ally the Australian Achras was added to the terminal to clarify the position of Metanoeus. clade which is characterised by straight slender costae Zoogeography (char. 27). In both cladograms (Figs 1, 2) Synchonnus is a basal member of the Hemiconderina. Although genera Unfortunately, the very fragmentary knowledge of classified in the Hemiconderina have many similar char­ Lycidae at the alpha taxonomic level and consequently acters, it can not be defined as a monophyletic group, as the doubtful information on generic distribution limit the the Hemiconderina clade has no autapomorphic character following discussion. I have used unpublished informa­ state. tion gathered by studying collections in the above institu­ That the Metriorrhynchina is monophylic is not well tions. The data in the lycid catalogue (Kleine, 1933) are founded, as I have not found any synapomorphic char­ inapplicable. The typological species concept applied to acter state to support the cladistic definition. In addition, most genera by Kleine and the problems originating from the relationships between the individual clades within the Pic’s confused work means that these data has to be con­ Metriorrhynchina remain obscure. firmed by studying the original type material. Information Most members of this clade have a complex pattern of on the distribution of genera is given in the systematic pronotal carinae forming four areoles on the frontal part. margin of the pronotum (i. e., frontolateral costae are pre­ On the other hand, Lycidae appear incapable of sent on the pronotum, character 18). This character is crossing natural barriers such as sea straits or deserts. useful for identifying most Metriorrhynchina. On the cla- Lycidae are often limited to humid mountain habitats and dogram in Fig. 1 the Metriorrhynchina without basal when these mountains are isolated, Lycidae easily speci- clade Bulenides and Caenioxylobanus share the state ate. Unlike many other groups, there are very few lycid species with a very wide range. The few widely distrib-

328 Figs 131-146: 131-132. Male genitalia, Mimoxylobanus reticulatus; 133-140. Ovipositor. 133 - Xylobanellus erythropterus; 134 - Metriorrhynchus sp.; 135 - Caenioxylobanus ater; 136 - Metriorrhynchoides sp.; 137 - Strophicus nigellus; 138 -Synchonnus appositus; 139 - Pseudodontocerus pulcher; 140 - Cladophorus sp.; 141-142. Female genitalia. 141 - Caenioxylobanus ater; 142 - Xylobanellus erythropterus; 143 - Pseudodontocerus pulcher, 144 Cladophorus sp.; 145-146 - Synchonnus appositus. Scales 0.5 mm. uted species are usually from lowlands and in this case Russian Far East through Siberia to Belarussia and the size of range is not impressive. I know only of one Poland (Fig. 3). The highest species and generic diversity species that occurs in two zoogeographical regions. occurs in the humid tropics of the Old World, but the The substantial part of the Metriorrhynchinae is an east generic species diversity is low in the Afrotropical Gondwanan element (distribution of Conderini is dis­ Region. There are also substantial differences in the con­ cussed further) with some representatives distributed in tribution of Metriorrhynchinae to the total representation East Asia and a single species widely distributed from of Lycidae in the zoogeographic regions.

329 The striking observation is that Conderini as a basal ippines (Bocak, 2000a) and only one species from the group within the Metriorrhynchinae have the distribution Asian continent and the Great Sundas. The distribution of similar to Erotinae, although they are not recorded from Microtrichalus is similar. Although its range reaches fur­ the Nearctic Region. Most species are known from the ther to the north and recently one species was found in south-eastern part of the Palaearctic Region and the southern Yunnan and northern Laos, the diversity of the northern part of the Oriental region, fewer species occur Oriental fauna is very low and the number of species distributed in the north-eastern part of the Palaearctic. decreases gradually from the Wallace line to the north­ Unlike the Erotinae they are not known from the Afro- west. Congruent with patterns found in other groups and tropical andNearctic Regions (Fig. 3). the tectonic history, Microtrichalus is rich in species in The range of Metriorrhynchini is limited to the Afro- the Philippines (Bocak, 1998d). tropical, Oriental, Australian and eastern part of The range of Metriorrhynchina is almost coincident Palaearctic Regions. It seems reasonable to assume that with that of the Metriorrhynchinae (Fig. 3). The the Metriorrhynchini dispersed to the Palaearctic Region Palaearctic fauna of Metriorrhynchina is poor, both in after they evolved somewhere on the Gondwana number of species and genera. Only Cautires and Xylo- continent. There are no endemic genera in the Palaearctic, banus are reported from China, Japan and the Russian Far and generally both the species and generic diversity of East. Metriorrhynchina are quite rare in the Palaearctic Palaearctic fauna are extremely low and decreases with Region and form a small part of the Palaearctic lycid the distance from the Oriental Region. Additionally, the fauna considering the number of species. Dispersal from Metriorrhynchinae form usually a small fragment of the the south offers a possible explanation for the present dis­ Lycidae collected in the Palaearctic Region, unlike in the tribution ofMetriorrhynchina in the Palaearctic Region. Oriental Region. The distribution of the Metriorrhynchini The Afrotropical fauna of Metriorrhynchina has not yet is given in Fig. 3. been revised and according to Kleine (1933) 192 species Another important observation is that within the Metri­ are reported from the Afrotropical Region. These species orrhynchinae the two Australian lineages, Hemiconderina are classified in genera Cladophorus, Cautires, Procau- and Trichalina, have a basal position and that the diver­ tires, Caenioxylobanus, and Xylobanus. I have studied sity of Australian fauna is much higher than Afrotropical available material in Pic’s and Kleine’s collections and I one. The Hemiconderina are endemic to the Australian found that the Afrotropical species placed in Cladophorus Region (Fig. 5) and within the subtribe the basal lineage and Procautires are not congeneric with Australian repre­ represented by Synchonnus is restricted to continental sentatives of these genera and should be placed in Cau­ Australia as well as Achras, which also has several plesi- tires. Afrotropical species of Stadenus were transferred to omorphic characters. The remaining genera of Hemicon­ the Erotinae (Bocak & Bocakova, 1992). Therefore, only derina are recorded from New Guinea and the Indonesian genera Cautires (about 150 species), Xylobanus (about 40 Archipelago east of the Wallace line. Both, species), and Caenioxylobanus (2 species) are now con­ Hemiconderis, known from New Guinea and the Yapen firmed for the Afrotropical Region. The Afrotropical Island, and Wakarumbia, from Sulawesi and the Buton fauna seems species rich, but multiple synonyms can be Island, have undergone considerable speciation. At pre­ expected as Pic often described common species several sent, 23 Wakarumbia species are recorded from Sulawesi. times. Caenioxylobanus reported from Madagascar is the Although, they show three different colour patterns, the only genus endemic to the region. substantial differences are in the male genitalia. Consid­ The Afrotropical and Indian faunas of Metrior­ ering the tectonic history of Sulawesi and the limited rhynchina are similar regarding the representation of gen­ range of Wakarumbia, we can presume that a part of era. Cautires, Xylobanus, and Prometaneus are the only Sulawesi of Australian origin or a part having contact genera occuring west of the low Brahmaputra River, and with some part forming today New Guinea brought the Prometanoeus is the only genus endemic to the western ancestor of Wakarumbia to Sulawesi. I have not found part of the Oriental Region. Cautires is a widely distrib­ any group of species restricted to one part of an island uted genus known from West Africa to the Philippines (Bocak, 1999b, 2000c, 2001b), therefore the simultane­ (Fig. 4) and Japan. Xylobanus has a similar distribution, ous speciation on various mountain ranges of widespread but unlike Cautires, Xylobanus crosses the Wallace line lowland species is possible. to the east. This genus has not yet been revised and the The Trichalina are the other basal group of presumably real relationship of the Australian representatives is Australian origin. All genera constituting the Trichalina unknown. are known from the Australian Region, only some of Metriorrhynchina from the eastern part of the Oriental them, Diatrichalus and Microtrichalus are present also in Region are more diverse. In addition to Cautires and the Oriental Region (Fig. 5). The highest diversity is gen­ Xylobanus, two endemic genera are common in South erally found in tropical parts of the Australian Region. East Asia - Metanoeus and Bulenides. Both do not cross After the revision of the Papuan fauna almost forty the Wallace line (Fig. 4). Metriorrhynchus is the only Diatrichalus species are recorded from the Australian genus in the Metriorrhynchina which has its centre of dis­ Region (Bocak, 2001a), and further species can be tribution in the Australian Region and reaches continental expected from Australia. On the other hand, only five Asia. Although this genus is reported from India (Kleine, Diatrichalus species were recently reported from the Phil­

330 Figs 147-163: 147-160. Female genitalia. 147 - Metriorrhynchoides sp.; 148 - Metriorrhynchus mirabilis; 149 - Cautires sp.; 150 - Metanoeus sp.; 151 - Achras limbatus; 152 - Pseudodontocerus sp.; 153 - Wakarumbia gracilis; 154 - Xylobanus costifer; 155 - Bulenides obsoletus; 156 - Stadenus puncticollis; 157 - Leptotrichalus accomodatus; 158 - Broxylus fenestratus; 159 - Sta­ denus triareolatus; 160 - Cautiromimus reticulatus. 161 - trochanter and femur. Pseudodontocerus pulcher; 162-163. Female sternum A7. 162 - Procautires sp.; 163 - Cautires sp. Scales 0.5 mm.

331 1933), it was not found in collections from localities west Trichalina with a basal placement, form the Metrior­ ofBurma. rhynchini. The Hemiconderina concept is redefined. There are several endemic lineages of Metriorrhynchina The genera Diatrichalus and Enylus are returned to the in the Australian Region: Pseudodontocerus, Stadenus, Trichalina. The resulting cladograms varied but some Ditua, Cladophorus, Cautiromimus, Broxylus, Kassemia, clusters within the Metriorrhynchina were relatively Porrostoma, and Metriorrhynchoides. The ranges of these robust (Ditua clade, Porrostoma clade and Cautires genera extend to the west in various degrees (Fig. 6). clade). Metriorrhynchus and Lobatang have most of their repre­ C. The distribution of clades was discussed. Conderini are sentatives in the Australian Region but a very small pro­ restricted to East and South East Asia, Metriorrhynchini portion of them cross the Wallace line onto the Asian are considered to be an East Gondwanan element with continent (Metriorrhynchus) or to the Philippines only a few species reaching northern East Asia. All (Lobatang). The distribution of Leptotrichalus is a bit dif­ these species are members of widespread genera known ferent. Most species of Leptotrichalus are reported from from the Afrotropical and Oriental regions. All major the Philippines, several from Sulawesi, the Lesser clades classified as subtribes of Metriorrhynchina are Sundas, Java, Sumatra and Borneo, and several from the either exclusively Australian or at least have most of Asian continent, only one of which occurs beyond the their endemic taxa in the Austro-Papuan Region. Isthmus of Kra (Bocak, 2000b). Recently, several unde­ D. The cladistic analysis showed that characters used for scribed species were collected in Halmahera. This sup­ a long time to define genera (presence and/or absence ports the conclusion about the relationship between of elytral secondary costae, shortened primary elytral Lobatang and Leptotrichalus inferred from morpho­ costae, and serrate of flabellate antennae) evolved sev­ logical data in phylogenetic analysis. Lobatang, is a eral times in the Metriorrhynchinae. closely related genus known from the Papuan Subregion, This study aimed to establish phylogenetic relationship Wallacea, and the Philippines. Most species are known between the genera of Metriorrhynchinae in order to from the eastern part of the region. Dispersal followed by study further individual clades within the subfamily. speciation of an ancestor inhabiting islands north of Aus­ More extensive sampling and new sources of data, espe­ tralia possibly lead to the present distribution and species cially on immature stages and molecular data, will cor­ richness of Leptotrichalus. roborate or refuse the proposed phylogeny. Additionally, Two clades are restricted to the Australian Region. The further studies are need to establish new combinations clade Porrostoma, Stadenus and Metriorrhynchoides is and to enlarge the data base available for cladistic analy­ the most common group in Australia with few species ses. known from New Guinea. The Ditua clade (node 27, Fig. 1) is recorded mainly from New Guinea and adjacent TAXONOMY islands. Few species occur in Australia. Metanoeus has an Metriorrhynchinae Kleine, 1926 ambiguous position in the present analysis and is the only Metriorrhynchinae Kleine, 1926a: 97. genus endemic to the Oriental Region, but related to Type genus.Metriorrhynchus Gemminger et Harold, 1869. clades whose centre of distribution is the Australian Region. Cladophorini Kleine 1928: 222; Bocak & Bocakova, 1990a: There are substantial differences in the relative repre­ 641. Type genus.Cladophorus Guerin-Meneville, 1830: Plate II, fig. sentation of Metriorrhynchinae within the Lycidae. Metri- 9. orrhynchinae are relatively poorly represented in Afro- tropical and Palaearctic Regions, where they usually form Dilolycinae Kleine, 1926: 186; Bocak & Bocakova, 1990a: 641. 10-20 per cent of collected specimens. They are more Dilolycini: Kleine, 1933: 84. Typegenus.Dilolycus Kleine, 1926a: 186. common in humid areas of South East Asia where they make up slightly over 50 per cent of the specimens. Their Haplothoracinae Kleine, 1926a: 95 (nomen nudum); Bocak & relative abundance increases to the east. East of the Wal­ Bocakova, 1990a: 641. lace line the Lycidae are almost exclusively represented Type genus.Haplothorax Kleine, 1926a: 95 (nomen nudum). by the Metriorrhynchinae. The Metriorrhynchinae form Diagnosis.Body small to large (3-30 mm), mostly 95 per cent of the specimens collected in New Guinea and slender, flat (Figs 7-9), weakly sclerotized. Head small, a substantial part of the Australian lycid fauna. Metrior­ partly concealed by pronotum. Cranium mostly compact rhynchinae form 90 per cent of the lycid species recorded (Fig. 11), seldom rostrate (Fig. 10). Antennae from Australia. compressed, filiform, serrate or flabellate (Figs 29-51). Conclusion Mouthparts well developed, mandibles small, moderately curved (Figs 24-28), maxillary palpi 4-segmented (Figs A. All ingroup terminal taxa form a monophyletic group 15-17), labial palpi 3-segmented. Labium simple (Fig. characterised by the presence of a circular phallobase 14). Pronotum with carinae (Figs 57-86). Elytra with four (Figs 91-132) and an antennomere 2 much shorter than primary and five secondary longitudinal costae (e. g. Figs antennomere 3 (Figs 29-51). Further apomorphic char­ 52-56), sometimes secondary costae absent (Figs 56, 87, acter states ofMetriorrhynchinae are given in Table 3. 88, 90) or primary costae reduced. Longitudinal costae B. The present analysis supports the tribes Conderini and connected by transverse costae. Male genitalia with para- Metriorrhynchini. Three subtribes, Hemiconderina and

332 meres and retracted, membranous internal sac (Fig. 91) or Diagnosis.Body small to medium sized, most species parameres absent and internal sac sclerotized or exposed cinnabar red. Antennae filiform to slightly serrate (Fig. (Figs 92-132). Ovipositor with plate-like coxites, small 29). Pronotum with five areoles (Fig. 58), sometimes movable styli and sclerotized, rod-shaped valvifers (Figs median areola reduced in size (as in Fig. 57), narrow, 133-140). Vagina mostly membranous, seldom sclero­ connected with frontal and lateral margins by stout cari­ tized (Figs 141-160). Spiculum gastrale moderately long nae. Elytra parallel-sided to slightly widened posteriorly, to absent (Figs 162-163). each elytron with four stout, longitudinal primary costae Distribution. Afrotropical, Oriental, and Australian Regions, and five much weaker, sometimes interrupted, secondary eastern part of Palaearctic Region, Siberia, Eastern Europe (Fig. costae. Transverse costae dense, sometimes irregular, 3). bottom of elytral areoles bald. Terminal male abdominal sternite with hole in basal part. Phallus almost straight in Remarks. Metriorrhynchinae are easily recognisable middle, parameres long (Fig. 91). Valvifers with attached within the Lycidae as they have pronotal carinae (Figs sclerites, widened in middle part, basal margin of coxites 57-86) and a very short antennomere 2 (Figs 30-51). In strengthened(Fig. 133). Vaginamembranous. addition all representatives have a circular phallobase (Figs 91-132), and parameres that are characteristically Distribution. Eastern part of the Palaearctic Region (China, slender or absent (Figs 91-132). Japan), Oriental Region (mainly continental South East Asia, less common in Himalayas and India, as well as in Sundas; east­ Conderini Bocak et Bocakova, 1990 ernmost record from Lombok). Over 20 species known from the Conderini Bocak et Bocakova 1990a: 643. range ofthe genus. Type genus.Conderis C. O. Waterhouse, 1879: 59. Remarks. Many Conderis species share the same char­ Diagnosis.Body small to medium-sized, most species acteristic elytral structure. The transverse costae are usu­ cinnabar red. Rostrum absent, antennae filiform to weakly ally very dense and the bottoms of elytral areoles are serrate (Fig. 29), almost circular in cross-section to bald, mat and the surface has a fungus-like structure. This weakly compressed. Pronotum with five areoles, median is the only apomorphic character of Conderis but not all areola often reduced in size, connected with pronotal and species. basal margins by long carinae (Figs 57, 58). Elytra with Xylobanellus Kleine, 1930 four strong primary costae, secondary costae present Xylobanellus Kleine, 1930a: 171. (Conderis) or absent (Xylobanellus, Fig. 87). Male geni­ Type species. Xylobanellus atricolor Kleine, 1930a: 171 (by talia with long parameres, base of phallus curved, phallo­ original designation). base with process at base, bases of parameres fused with phallus to form a unique process (Fig. 91). Ovipositor Rossioptera Kasantsev, 1988: 169,syn. n. Type species.Eros erythropterus Baudi, 1871 (by monotypy). robust, with stout valvifers, small sclerites attached to val­ vifers (Fig. 133). Vagina membranous, unpaired gland Chuzenjianus Kono, 1932: 61; Nakane, 1969a: 176. very small, inconspicuous, lateral glands very delicate Type species. Chuzenjianus tenuis Kono, 1932: 61 (by original and very wide at base (Fig. 142). designation). Distribution. Palaearctic and Oriental Regions. Over 30 spe­ Diagnosis.Body small to medium-sized, slender, cies described at present; mainly from China and continental parallel-sided. Antennae slender, long, slightly serrate. South East Asia. Apical maxillary palpomere triangular. Pronotum almost parallel-sided, posterior angles rectangular. Median areola Remarks. Two genera, Conderis and Xylobanellus are small, very slender (Fig. 57). Elytra flat, with four stout classified as Conderini. Their close relationship is well longitudinal costae, transverse costae well marked, only supported by the unique shape of male (Fig. 91) and 20-25 costae per interspace, elytral areoles squared to female genitalia (Figs 133, 142). They are easily distin­ longitudinal (Fig. 87), their bottom flat, matt, finely cha­ guished by the presence or absence of secondary elytral grined. Male genitalia with slender curved phallus, its costae. The absence of secondary costae is the only apex widened in lateral view. Parameres well developed. advanced feature found in Xylobanellus, but this occurs Ovipositor robust, vagina membranous. independently in many lineages within the Lycidae (sev­ eral occurences in the Metriorrhynchinae). Unfortunately, Distribution. Five species are placed in Xylobanellus, one Conderis does not have any synapomorphic character occurring in Eastern Europe and Siberia, the rest in China, Thai­ state and therefore Conderis can be paraphyletic with land, Malaysia, Borneo and Java. respect to Xylobanellus. Remarks. All Xylobanellus species share several char­ Conderis C. O. Waterhouse, 1879 acters, which potentially could be present in a common ancestor: secondary costae absent, rectangular reticulate Conderis C. O. Waterhouse, 1879: 59. cells and male segment A7 with straight margin. The Type species. Calopteron signicolle Kirsch, 1875: 36; C. O. Waterhouse, 1879: 59 (by original designation). similarly structured elytral costae is also encountered in unrelated lineages within the Metriorrhynchinae (e. g. Pseudoconderis Pic, 1921a: 8, hors texte; Bocak, 1998a: 18. Broxylus, Caenioxylobanus, Xylobanus) and in other sub­ Type species. Pseudoconderis gorhami Pic, 1921a: 8; Bocak, 1998a: 18 (by subsequent designation). families of Lycidae (Lycinae: Calopterini, part; Erotinae, part). Rossioptera is based on Eros erythropterus, previ­ ously classified in Dictyoptera. Bocak & Bocakova

333 (1987a) transferred E. erythropterus to Xylobanellus. straight carinae forming only five areoles on the pronotal Rossioptera is considered to be a junior subjective disc (Figs 59-60), head without rostrum, and primary synonym of Xylobanellus. costae 1 and 3 weaker apically; rarely also a costa 1 partly Metriorrhynchini Kleine, 1926 shortened (Hemiconderis bipustulatus Bocak & Bocak­ ova, 1990b). Additionally, all genera placed here share a Metriorrhynchini Kleine, 1926a: 97. slender, delicate appearance, with parallel-sided elytra, Type genus. Metriorrhynchus Gemminger et Harold, 1869: 1629. very slender legs, slender and relatively long maxillary palpi, slightly securifom apical maxillary palpomere, Diagnosis.Body small to large. Head mostly lack a ros­ minute labial palpi, and absence of tibial spurs. trum, seldom has a more or less long rostrum. Antennae serrate to flabellate in males, serrate in females, seldom Synchonnus C. O. Waterhouse, 1879 flabellate in females. Pronotum with only median areola Synchonnus C. O. Waterhouse, 1879: 59. and lateral costae present (Figs 61-65, 84), with slender Type species. Synchonnus clientulus C. O. Waterhouse, 1879: costae forming five areoles (Figs 59-60) or with seven 59 (by monotypy). areoles (e. g. Figs 76-83). Elytra with longitudinal and Diagnosis.Body small, quite robust, slightly widened transverse costae. Secondary costae weak to absent (Figs posteriorly. Head small, without rostrum. Antennae 52-56, 87-90). Male genitalia without parameres (Figs robust, strongly compressed, serrate. Maxillary palpi 92-132), with partly sclerotized internal sac, or large, slender, apical palpomere slightly widened. Pronotum usually finely setose sac exposed on apical part of phallus with strongly projected hind angles, with five areoles, (Fig. 118). Vagina with paired, lateral, narrowly attached median areola slender, long. Four primary and five glands (Figs 143-160), and with apparent, flat, unpaired secondary costae extant whole length of the elytra, gland (e. g. Figs 143, 145). primary costa 2 and 4 much stronger, costa 3 very weak Distribution. Afrotropical, Oriental, and Australian Regions apically. Secondary costae weak, transverse costae and eastern part ofPalaearctic Region (Fig. 3). irregular. Male genitalia very slender, phallus widened in Remarks. Metriorrhynchini are easily recognisable by apical third, internal sac with two large spines in basal the absence of parameres, presence of characteristic pro­ part. Ovipositor with small coxites and long valvifers notal carinae forming up to seven areoles, flat unpaired (Fig. 138), vaginawithdorsal sac (Figs 145-146). vaginal gland and narrowly attached lateral glands. Metri­ Distribution. Australian Region. Five species reported from orrhynchus Guérin-Méneville, 1838 was considered to be Australia, New Guinea, and Jobi Island. the type genus by Kleine (1926a). Bocak (1998c) dis­ Remarks. Synchonnus has a basal position in cussed the validity of the name. Hemiconderina. As in Achras the male genitalia has a pair Hemiconderina Bocak et Bocakova, 1990 of spines (Figs 92-93, 95-96), but unlike this genus it has Hemiconderina Bocak et Bocakova 1990a: 645. more robust body and obtuse pronotal carinae. Typegenus.Hemiconderis Kleine, 1926a: 162. Achras C. O. Waterhouse, 1879 Diagnosis. Body small to medium-sized, mostly Achras C. O. Waterhouse, 1879: 61. slender, parallel-sided (Wakarumbia, Hemiconderis, Fig. Type species.Achras limbatus C. O. Waterhouse, 1879: 62 (by 8, and Achras), seldom shorter, more robust monotypy). (Synchonnus). Head small, without rostrum. Antennae Diagnosis.Body medium-sized. Head without rostrum. serrate (Figs 30, 31), antennomeres triangular to parallel­ Antennae strongly compressed, slender, only sided, always apparently compressed, similar in both antennomeres 3 and 4 broad, subsequent antennomeres sexes. Pronotum with five areoles, carinae straight, slen­ narrow, acutely pointed. Maxillary palpi quite robust. der, median areola small, narrow (Figs 59, 60). Elytra Pronotum with almost straight basal margin and posterior parallel-sided to slightly widened posteriorly (Fig. 8). angles acutely projected. Disc with five areoles. Elytra Elytral costae 2 and 4 stronger, always reaching apex of with strong primary costae on humeral third of elytra, elytra, costae 1 and 3 always apparently weaker in apical much weaker further back, primary costae 1 and 3 much half of elytra, seldom shortened. Secondary costae present weaker apically, secondary costae on apical third of costa in most genera, absent in Wakarumbia. Legs slender, 3 absent, on costa 1 irregular. Transverse costae on elytra strongly compressed. Male genitalia with simple, slender weak, irregular, elytral cells mostly slightly transverse. phallus (Figs 92-96). Internal sac with pair of basal Male genitalia robust, short, with two large spines on spines, seldom fused in one complex structure (Wakarum­ basal part of internal sac. Ovipositor with short coxites bia, Fig. 94). Ovipositor wide, valvifers apparently diver­ and long, slender valvifers, vagina large, membranous gent, vaginamembranous, simple (Figs 151, 153). (Fig. 151). Distribution. Australian Region: Australia, New Guinea, Distribution. Australian Region: Australia (New South Yapen Island, Sulawesi, Buton Island (Fig. 5); there was no Wales). Monotypic. overlap in the distribution of Wakarumbia, Achras, and Hemi- conderis. Remarks. Achras was classified with Metriorrhynchus and Metriorrhynchoides in the Metriorrhynchini (Kleine, Remarks. Hemiconderina are easily distinguished by 1933) and later was not studied but kept in the Metrior- the shape of their pronotum, with characteristic, slender, rhynchina (Bocak & Bocakova, 1990a). Based on the pre­

334 sent analysis it is transferred to the subtribe Hemicon- most species apical part of phallus rotated to various derina. Achras is characterised by sclerotized, widened degrees. basal part of spermaduct and lateral sacks at base of Distribution. Australian Region: Sulawesi, Buton Island. vagina. The second character can very between species Altogether 23 species classified in Wakarumbia. analogous to the situation in other genera. Achras seems Remarks. Wakarumbia is the only genus of Hemicon­ to be very closely related to Wakarumbia but retains sev­ derina that lacks secondary costae. Apical part of phallus eral plesiomorphic character states, i. e. vestiges of secon­ in most species is characteristically rotated and there is dary costae, unrotated phallus, pair of basal spines on internal sac. one complex spine at base of the phallus. Wakarumbia and Achras both have characteristically Trichalina Kleine, 1928 shaped valvifers (Fig. 151), vagina with unpaired gland Trichalinae Kleine, 1928: 222. located on independent frontal sac, both genera have Trichalini Kleine 1933: 69. much weaker costae 1 and 3, but which extent the whole Trichalina Bocak et Bocakova, 1990a: 646 length of elytra. They have similar, characteristic labrum Type genus.Trichalus C. O. Waterhouse, 1877: 82. and slender lateral processes on mesoscutum. Diagnosis.Body small to medium-sized. Head without Hemiconderis Kleine, 1926 rostrum. Antennae serrate, seldom with short lamellae Hemiconderis Kleine, 1926a: 162. (several Diatrichalus, Fig. 32), or flabellate (Flabellotri- Type species.H. explicatus Kleine, 1926a: 162 (by monotypy). chalus). Pronotum with slender median areola (Figs Diagnosis.Body small to medium-sized, slender, col­ 62-65), with two divergent carinae (Eniclases, Fig. 66) or oration usually bright, seldom dull. Head small, without with five areoles (Schizotrichalus). Elytra regularly with rostrum. Antennae considerably compressed, weakly ser­ considerably shortened primary costa 1 (Fig. 9), secon­ rate, reaching beyond elytral apex. Maxillary palpi rela­ dary costae usually well developed, absent in some tively long, slender, terminal palpomere slightly widened. Diatrichalus. Male genitalia slender with basal spines Pronotum with slender, conspicuous carinae forming 5 (Figs 97-99), widened apically with complex sclerotized areoles. Median areola slender (Fig. 60). Elytra long, internal sac (Figs 101-102) or slender, with partly sclero­ slender, with 4 primary costae at base, primary costa 3 tized apical part, internal sac without basal spines (Figs shortened, reaching middle of elytra. Primary costa 1 104-106) or basal spines apparent (Fig. 103). Ovipositor seldom short. Secondary costae well developed, reticulate with simple coxites, valvifers usually long and slender, cells small, regular. Phallus lanceolate, internal sac with vagina always membranous, lateral glands usually dor­ two large spines linearly arranged. Female genitalia sally attached (Microtrichalus etc.), sometimes laterally broad, well sclerotized, coxites free. Valvifers thin, their attached (Diatrichalus, Enylus only). bases with various projections, widened. Distribution. Australian Region and eastern part of the Ori­ ental Region: Indochina, Thailand, southernmost Yunnan, Pen­ Distribution. Australian Region: New Guinea, Yapen Island. insular Malaysia, the Philippines, and the Great Sundas (Fig. 5). Altogether 9 species known (Bocak & Bocakova, 1990b). Highest diversity in New Guinea and Australia, the number of Remarks. Hemiconderis is the only member of the species decreases to the west of the range and only a few species Hemiconderina having entire secondary costae, shortened are known from continental Asia. primary costa 3 and two spines on internal sac linearly Remarks. The genera with shortened elytral costa 1 arranged; the first located at base of phallus, the second previously classified in the tribe Trichalina were revised on apical half. Basal part of valvifers of all females wid­ recently by Bocak (1998b). Only genera with the charac­ ened to some degree (Bocak & Bocakova, 1990b). H. teristic slender phallus and dorsally attached lateral bipustulatus Bocak & Bocakova, 1988 has additionally a vaginal glands belonged in the Trichalina in the restricted shortened costa 1, but to a considerably smaller degree sense, and the sister-group position of Diatrichalus and than encountered in Trichalina. Enylus was confirmed. The present analysis of the whole Wakarumbia Bocak, 1999 subfamily Metriorrhynchinae confirmed the monophyly Wakarumbia Bocak, 1999b: 166. of the restricted Trichalina and also supports the close Type species. Wakarumbia gracilis Bocak, 1999b: 167 (by relationship of genera Enylus and Diatrichalus to the Tri­ original designation). chalina clade. Although the relationship of these genera needs confirming using larval characters and molecular Diagnosis.Body small, very slender, delicate, with data, the genera Enylus and Diatrichalus are here classi­ slender legs and antennae. Head without rostrum. fied in the Trichalina. Antennae slightly serrate to parallel-sided. Pronotum with five areoles, pronotal carinae straight and slender. Elytra Enylus C. O. Waterhouse, 1879 almost parallel-sided, with four primary costae, primary Enylus C. O. Waterhouse, 1879: 72. costae 1 and 3 weak, secondary costae absent, only ves­ Type species.E. segregatus C. O. Waterhouse, 1879 (by mono- tiges present at base of elytra. Transverse costae well typy). developed, regular, elytral cells mostly square-shaped. Strophicus C. O. Waterhouse, 1879: 73,syn. n. Phallus with one, complex basal spine on internal sac, in Type species. Strophicus nigellus C. O. Waterhouse, 1879: 73 (by monotypy).

335 Diagnosis.Body small to medium-sized. Head small, inserted laterally. Spermaduct robust at base, spermatheca without rostrum. Antennae strongly compressed, similar slender, very long. in both sexes, serrate or antennomeres almost parallel­ Distribution. Australian and eastern part of the Oriental sided, never flabellate. Maxillary palpi robust, apical pal- Region. Only one species found in continental Malaysia, Java, pomere with an oblique apex, broad. Pronotum only with Borneo and Sumatra. All other Oriental representatives occur in median areola, lateral carinae absent, weak lateral tuber­ the Philippines. The highest diversity is found in New Guinea. cles on lateral margins of pronotum. Elytra with four pri­ Kleine (1933) placed six species inDiatrichalus. Later, it was mary costae, primary costa 1 strong over whole length, found that many Oriental and Papuan species previously costae 2 and 4 much stouter than costae 1 and 3, costa 3 classified as Trichalus belonged to Diatrichalus (Bocak, 2000a, much weaker apically. Legs slender, strongly 2001a). At present over 40 species are classified in this genus. compressed. Male genitalia always with slender, long Remarks. In all Diatrichalus the basal part of phallus, with pair of basal spines, partly sclerotized api­ spermaduct is widened and this unique character is cally, phallobasal membrane inconspicuous. Ovipositor considered to be apomorphic state. This genus originally with short, rather stout valvifers, fused with coxites (Fig. included only species that lacked secondary costae. A 137), vagina relatively short, bases of glandular ducts study of most species showed that in Diatrichalus there robust, accessory glands inserted laterally. Spermatheca are transitional states between strong secondary costae, slender, very long. weak irregular secondary costae and complete absence of Distribution. Australian Region: New South Wales, New costae. The internal sac and basal part of female Guinea, and Mysool Island. Only four species are at present spermaduct in Diatrichalus has a very characteristic and classified in Enylus, two of them were originally classified in unique shape of and both species that have and lack Strophicus. secondary costae are included in this genus. Remarks. Enylus is the only genus classified in Tri- Trichalus C. O. Waterhouse, 1877 chalina not having a shortened elytral primary costa 1. Trichalus C. O. Waterhouse, 1877: 82. Only the holotype of Strophicus nigellus was available, Type species. T. flavopictus C. O. Waterhouse, 1877: 82; C. O. and therefore Strophicus was not included in the phyloge­ Waterhouse 1878: 103 (bysubsequentdesignation) netic analysis. Strophicus is externally very similar to Xantheros Fairmaire, 1877: 167 Enylus and the only differs in the size of its eyes: very Type species. X. ochreatus Fairmaire, 1877: 167; Bourgeois small in males of Strophicus and very large in Enylus. 1891: 347 (by subsequentdesignation). The presence and/or absence of sexual dimorphism in eye Diagnosis. Body medium-sized, slightly widened size ise regularly observed in genera of Lycidae. This posteriorly, antennae serrate in both sexes. Pronotum with character is correlated with when in the day they fly. one longitudinal median areola, first elytral primary costa Males of species active at night have large eyes. There­ shortened, secondary costae always well developed, fore, I propose that Strophicus is a junior subjective regular. Male genitalia with slender phallus, internal sac synonym of Enylus. membranous, with pair of spines at base, internal sac Diatrichalus Kleine, 1926 exposed, free. Ovipositor with extremely robust valvifers, Diatrichalus Kleine, 1926a: 167. fused in basal third, flat, closely attached to coxital Type species. D. xylobanoides Kleine, 1926 (by original desig­ processes. Vagina robust, lateral glands attached dorsally, nation). their bases widened, unpaired vaginal gland short, Mimotrichalus Pic, 1930: 92, hors texte; Bocak, 1998b. spermathecal duct short, spermatheca large, bulbous. Type species.M. tenimberensis Pic, 1930 (by monotypy). Distribution.Australian Region: only continental Australia. Diagnosis.Body medium-sized, coloration variable but Remarks. Kleine (1933) cited 112 Trichalus species mostly black or black with metallic blue shine, only a few from the Oriental and Australian Regions. Recent studies species have a brightly coloured pronotum and basal part indicate that most Oriental and Papuan species previously of elytra. Never light concolour. Head small, without ros­ classified as Trichalus belong to Microtrichalus (Bocak, trum. Antennae strongly compressed, antennomeres 1998b, d, 2000d). The AustralianTrichalus is badly in parallel-sided to acutely serrate in both sexes, seldom need of revision. Widened bases of lateral glands and very shortly flabellate in male. Maxillary palpi with broad fused valvifers are the unique apomorphic character states apical palpomere bearing papillae at apex. Pronotum with of Trichalus. one more or less broad longitudinal areola in the middle, Microtrichalus Pic, 1921 lateral margins elevated. Elytral primary costa 1 consid­ erably shortened, reaching at most one third of elytral Microtrichalus Pic, 1921b: 9, hors texte. length, secondary costae irregular to completely absent in Type species.M. singularis Pic, 1921b: 9 (bymonotypy). some species. Legs relatively strong and short, strongly Falsoenylus Pic, 1926: 29, hors texte; Bocak, 1998b. compressed. Male genitalia with relatively short and Type species.F. basipennis Pic, 1926: 30; (by monotypy). robust phallus, internal sac lacks basal spines, regularly Diagnosis.Body small to medium-sized, very often strongly sclerotized apically (Figs 100-101). Ovipositor with dorsal part of body at least partly yellow to lightly with slender, long valvifers, sometimes incompletely brown. Antennae serrate in both sexes, labial palpi slen­ fused at base, vagina relatively short, vaginal glands der, apical palpomere slightly narrowed at apex, corn-

336 pressed. Pronotum with one longitudinal median areola. Type species. T. nigrescens C. O. Waterhouse, 1879: 70 (by First elytral primary costa shortened, secondary costae original designation). regularly present. Male genitalia very uniform within Diagnosis.Body small to medium sized, dark brown to genus. Phallus slender, only basally completely sclero- black. Antennae slightly serrate in both sexes. Maxillary tized, ventrally open apically, internal sac exposed, with palpi slender. Pronotum with five areoles. Elytral pri­ pair of sickle-like thorns at base, sometimes with mary costa 1 shortened, usually secondary costae devel­ V-shaped rod in membrane. Whole of male genitalia oped, in one (presently undescribed) species irregular often very slightly pigmented and sclerotized, rendering transverse costae and mostly absent secondary costae. structure of internal sac inconspicuous. Ovipositor with Male genitalia with long, slender, only basally well slender valvifers. Vagina slender to moderately robust, sclerotized phallus, dorsally one strengthened pigmented with dorsally attached lateral accessory glands whose keel, bearing reversed cup-like apex (Fig. 105). Middle bases are more sclerotized, proper glands very fine. Two part membranous, with internal sac pigmented apically, lateral pockets of unknown function are present in lateral no basal spines present. Phallobase without extensive part of vagina. Very slender unpaired gland attached to membrane. Ovipositor with long slender valvifers, some­ base of vagina. times connected at bases. Vagina long, slender, lateral Distribution. Australian and Oriental Regions: Sunda accessory glands inserted dorsally, apical gland short. Islands, the Philippines, Peninsular Malaysia, Thailand, Indo­ Spermaduct short, straight, spermatheca robust. china, and the southernmost part ofYunnan. Distribution. Australian Region: New Guinea. Monotypic, Remarks. Many species formerly placed in Trichalus but several undescribed species have been collected. were transferred to Microtrichalus (Bocak, 1998d, 2000 d). At present, 40 species are classified in Microtrichalus, Remarks. Schizotrichalus differs in five pronotal are­ but the faunas of New Guinea and Australia, where the oles from other genera classified in the in Trichalina. highest diversity is found, have not yet been revised. Lat­ Eniclases C. O. Waterhouse, 1879 eral pockets and unpaired basal vaginal gland are the Eniclases C. O. Waterhouse, 1879: 66. unique apomorphic character states of Microtrichalus. Type species. Lycus (genus 35) luteolus C. O. Waterhouse, Flabellotrichalus Pic, 1921 1878 (by original designation). Flabellotrichalus Pic, 1921b: 9, hors texte. Trichalolus Pic, 1923: 36, hors texte; Bocak & Bocakova, 1991. Type species.F. notatithorax Pic, 1921b: 9; Kleine, 1936: 4 (by Type species. T. apertus Pic, 1923: 36 (bymonotypy). subsequent designation). Diagnosis.Body medium-sized. Head without rostrum, Stereotrichalus Kleine, 1926a: 183; Kleine, 1930a: 330. antennae with parallel-sided to serrate antennomeres Type species. S. evidens Kleine, 1926a: 184 (by subsequent 3-10, only a few species antennomeres flabellate in male. designation) Labial palpi slender, apical palpomere parallel-sided. Pro­ Villosotrichalus Pic, 1921b: 9, hors texte; Bocak, 1998. notum divided by two divergent carinae into three fields Type species. V.reductus Pic, 1921b: 9 (by monotypy). (Fig. 66). Elytra with first elytral primary costa shortened, with complete secondary costae. Male genitalia with Diagnosis.Body medium sized to large, slightly wid­ ened posteriorly. Usually dark brown to black with apparent keel on dorsal surface of phallus, with free, humeral part of elytra lighter. Head without rostrum, membranous internal sac, without sclerotised structures or spines (Fig. 106). Ovipositor with basal processes of cox­ antennae flabellate in male (Fig. 27). Pronotum with one longitudinal median areola, frontal and lateral margins of ites attached to valvifers, lateral glands of vagina attached pronotum very often with dense short to very long pubes­ into the middle part of vagina, glands very fine. cence. Elytral primary costa 1 shortened, secondary Distribution. Australian Region. At present, 26 species are costae always present. Male genitalia very slender, phal- known from New Guinea and one from the Moluccas. Although lobase usually covered with extensive membrane. Phallus I have seen very extensive lycid material from the Solomon very long, narrow, sclerotized completely only basally, Islands it contained no species ofthis genus. apically sclerotized dorsally, internal sac exposed, pig­ Remarks. Eniclases is characterised by a unique mented at apex, sometimes with strengthened rod in arrangement of pronotal costae. Bocak & Bocakova membrane, never with pair of spines at base. Vagina with (1991) revised this genus. two dorsally attached accessory glands, glands very fine, Metriorrhynchina Kleine, 1926 spermaduct long, curved, spermatheca lemon-shaped. Metriorrhynchina Kleine, 1926a: 97; Bocak & Bocakova, Distribution. Australian Region. Flabellotrichalus is 1990a: 644. recorded only from New Guinea and Moluccas. At present 13 Type genus. Metriorrhynchus Gemminger et Harold, 1869: species are classified in this genus. 1629. Remarks. Long antennal lamellae in male, pubescence Diagnosis.Body small to large. Head usually without on pronotum, and shape of male genitalia are unique rostrum, several genera with a very small (Leptotrichalus) characters of Flabellotrichalus within the Trichalina. or long rostrum (Porrostoma, Fig. 10, Metriorrhynchoi- Schizotrichalus Kleine, 1926 des, Oriomum). Male antennae often flabellate, the posi­ Schizotrichalus Kleine. 1926a: 183. tion of lamellae or antennomeres variable within genera,

337 lamellae usually robust (Fig. 38), seldom extremely Type species. Caenioxylobanus ater Pic, 1922 (hereby desig­ slender and long (Pseudodontocerus, Fig. 45, Cladopho- nated). rus, Figs 44, 51). Pronotum usually with distinct fronto- Diagnosis.Body medium-sized, completely dark brown lateral and lateral carinae and small, slender median to black. Head without rostrum. Antennae robust, areola attached to basal margin of pronotum (e. g. Figs strongly compressed, flabellate in male, serrate in female 69-74, 76-83), several genera with either only median (Fig. 39). Maxillary palpi robust. Pronotum trapezoidal, areola present and lateral costae vestigial (Leptotrichalus, with a distinct protruding frontal margin, which is formed Lobatang, Fig. 75), only frontal keel and median areola by a robust vertical surface (Fig. 67). One longitudinal strongly marked and lateral and frontolateral carinae ves­ keel present on disc, anterior half of keel sharp and nar­ tigial (Bulenides, Fig. 68), or only frontal keel distinct row, higher than wide at base, in basal part almost absent, (Caenioxylobanus, Fig. 67). Elytra with primary and sec­ transformed into a very wide, flat mound. Elytra flat, with ondary longitudinal costae connected by transverse costae four longitudinal costae and slender transverse costae. All (Figs 52-55, 89), secondary costae sometimes absent costae very regular, slender and sharp. Reticulate cells (Figs 56, 88, 89). Male genitalia without parameres, often quadrate to slightly transverse (Fig. 88), all costae shiny, lanceolate with pair of spines apically (Figs 107-12), black, bottom of cells velvet, dark brown. Male genitalia robust, with basal spiens (Figs 125-28) or tubular with slender, lanceolate, internal sac with two spines on apical exposed membranous internal sac (Figs 115-20). Female third. Ovipositor robust (Fig. 135), vagina membranous, genitalia with simple coxites, membranous or more or spermatheca long, bulbous apically (Fig. 140). less sclerotized vagina, large unpaired gland and pair of Distribution. Afrotropical Region: Madagascar. Two species lateral glands. are endemic to Madagascar. Distribution. Afrotropical, Oriental, Australian Regions and Remarks. Pronotal carinae of Caenioxylobanus have a eastern part of the Palaearctic Region. Coincident with distribu­ tion ofMetriorrhynchini (Fig. 3). unique shape and this genus differs from the closely related Bulenides in the absence of secondary costae and Bulenides C. O. Waterhouse, 1879 in the mat, velvet bottoms to the elytral cells. Bulenides C. O. Waterhouse, 1879: 34. Type species. Lycus (genus 21) obsoletus C. O. Waterhouse, Xylobanus C. O. Waterhouse, 1879 1878: 109; Bourgeois, 1891: 345 (bysubsequentdesignation). Xylobanus C. O. Waterhouse, 1879: 38. Type species. Lycus costifer Walker, 1858: 282 (by original Diagnosis.Body small to medium sized. Head without designation). rostrum. Antennae robust, flabellate in male, lamellae short and robust (as in Fig. 38), serrate in female. Maxil­ Invalid designation. Xylobanus fastidiosus C. O. Waterhouse, 1879: 39; Bourgeois, 1891: 345. lary palpi slender, apical palpomere pointed. Pronotum often narrowed anteriorly, with robust frontal margin Diagnosis.Body small to medium-sized, often slender, (Fig. 68), disc with one small median areola, regularly parallel-sided. Head without rostrum. Antennae serrate to with vestiges of frontolateral costae. Elytra with four flabellate in male, lamellae if present short, robust (as in strong, entire primary costae and weak but regular secon­ Fig. 38), more or less serrate in female. Maxillary palpi dary costae. Elytral cells minute. Phallus slender, lanceo­ robust, apical palpomere almost parallel-sided to consid­ late, internal sac with two spines on apical third of erably widened. Pronotum trapezoidal, usually with seven phallus. Vagina membranous, spermaduct short, sper- strongly marked areoles (Fig. 69), in some species lateral matheca lemon-shaped (Fig. 155). and frontolateral carinae obtuse to reduced. Elytra with four well developed, strong longitudinal primary costae, Distribution. Eastern part of Oriental Region: Burma, Thai­ land, Indochina, Taiwan, Peninsular Malaysia, Sumatra, Java, secondary costae absent, elytral reticulate cells transverse, Borneo, and the Philippines. regular. Male genitalia slender, phallus lanceolate as in Cautires (Fig. 108) or widened in apical part (Figs 109, Remarks. Bulenides is characteristic in having only a 110). Ovipositor short, with robust valvifers (Fig. 154), small, narrow areola in middle of pronotum, the carinae vagina short, membranous, spermatheca long, slender to forming the areola can be obtuse to absent and conse­ lemon-shaped. quently only a strong frontal keel is present on pronotum. The shape of pronotum is variable, but often distinctly Distribution. Afrotropical, Australian, and Oriental Regions, narrowed frontally. The shape of pronotum is similar to Palaearctic Region: China, Japan, Korea, and Russian Far East. Almost 300 species are described forthis genus. that of Caenioxylobanus, which differs in the complete absence of secondary elytral costae and tibial spurs. Addi­ Remarks. Xylobanus is an extremely species rich and tionally, the form of the frontal margin of the pronotum in widespread genus. It is characterised by a pronotum with Bulenides and Caenioxylobanus is unique (Figs 67-68). seven areoles (although the carinae can be obtuse to ves­ This character state was confirmed in B. obsoletus (type tigial on some parts of the pronotum, frontolateral and lat­ species) and some other species but is absent from some eral pronotal carinae are never completely absent) and species included in Bulenides. Only a complete revision elytra lack secondary costae (they are present at most ves­ of Bulenides will clarify the limit of the genus. tigially on narrow humeral part). Xylobanus and Cautires have a similar shaped pronotum and male and female Caenioxylobanus Pic, 1922 genitalia. Their close relationship is supported by their Caenioxylobanus Pic, 1922: 13, hors texte.

338 very similar larvae (Potozkaja, 1981, Bocak & Matsuda, have reliable synapomorphic character state for Xylo­ in press). The affinities with Broxylus are discussed fur­ banus, and therefore Broxylus may be a modified member ther. Several characters of Xylobanus are variable. The of Xylobanus. phallus is widely rounded apically and primary gonoporus Cautires C. O. Waterhouse, 1879 is sclerotized in many species, including the type species Cautires C. O. Waterhouse, 1879: 36. Xylobanus costifer, but the apex is pointed and the gono­ Type species. Lycus (gen. 22) excellens C.O. Waterhouse, porus membranous in X. corporaali Pic, 1922 from 1878: 110; Bourgeois, 1891: 345 (bysubsequentdesignation) Sumatra. On the other hand these species both have phallus with a wide base. Xylobanus costifer has valvifers Diagnosis.Body small to medium sized. Head without connected at base, as does one species from Bali and rostrum. Antennae flabellate in male (Fig. 38) and serrate some other species, but this sclerite is vestigial or absent in female. Maxillary palpi robust, apical palpomere wid­ in other species. Also presence of spiculum gastrale is ened, securiform. Pronotum trapezoidal, disc with seven variable and some species have flabellate antennae (e. g. areoles (Fig. 71), seldom lateral carinae partly reduced. X. nigrimembris Pic, 1926d), or serrate antennae (X. Elytra with four entire longitudinal primary costae. Sec­ humifer Walker, 1858). Only an extensive, detailed alpha- ondary costae considerably weaker, but well developed. taxonomical study can clarify the concept of Xylobanus. Transverse costae regular, as strong as secondary longitu­ The absence of secondary costae is an advanced feature, dinal costae. Phallus slender, lanceolate, internal sac with which has evolved several times in Lycidae. two spines on apical third of phallus (Fig. 108). Ovi­ positor slender, valvifers subtle, vagina membranous, Broxylus C. O. Waterhouse, 1879 spermatheca lemon-shaped (Fig. 149). Broxylus C. O. Waterhouse, 1879: 20. Distribution. Afrotropical and Oriental Regions; Palaearctic Type species. TCalopteron pfeifferi C. O. Waterhouse, 1878: Region: Russian Far East, China, Japan, Taiwan. Although Cau­ 106 (by monotypy). The name was cited by Waterhouse (1878) tires was reported by Kleine (1926a, 1933) from New Guinea, I with a question mark before the generic name indicating provi­ have not found any Cautires from this region and Kleine’s illus­ sional generic classification. trations of male genitalia are similar to those of Cladophorus or Samanga Pic, 1921b: 9, hors texte,syn. n. Cautiromimus. In the south-east, the furthermost records of Type species.Samangafenestrata Pic, 1921b: 9 (by monotypy). Cautires are from the Philippines, Borneo and Lesser Sundas Diagnosis.Body medium-sized. Head small, antennae (Sumbawa). I have not found any representative in Sulawesi or Moluccas and it is not present in New Guinean material. The reach beyond the middle of the elytra, compressed. Palpi highest diversity occurs in continental South East Asia, strongly compressed, maxillary palpi with long, spoon­ Sumatra, Java and Borneo. Cautires is the commonest lycid in like apical palpomere, palpomere 4 as long as combined Madagascar, but it is rare in continental Africa, although many length of palpomeres 1-3, labial palpi short, apical pal­ species are reported from there. pomere large (Fig. 15). Pronotum with partly reduced Remarks.All characters used for the definition of Cau­ carinae, frontolateral carinae present on areola only, lat­ tires (seven pronotal areoles, entire elytral costae and eral carinae weak (Fig. 70). Elytra flat, widest in apical shape of genitalia) are symplesiomorphic character states tenth or strongly hemispherically widened, with four lon­ and at present no apomorphic character state is known. gitudinal, sharp primary costae and thin, sharp transverse An alpha taxonomic study of the Cautires clade will lead costae of similar strength. Elytral cells quadrate to appar­ to a better understanding of the relationships. ently longitudinal (Fig. 90). Legs slender, strongly com­ pressed. Trochanters very slender, long. Male genitalia as Prometanoeus Kleine, 1925 figured (Fig. 111). Prometanoeus Kleine, 1925c: 133. Distribution. Australian Region: Sulawesi. Altogether 11 Type species. Prometanoeus ochraceus Kleine, 1925c: 133 (by species known from Sulawesi (Bocak & Jass, in press). monotypy). Tapromenoeus Bocak and Bocakova, 1989: 327,syn. n. Remarks. Broxylus was classified in the Calopterini Type species. Xylobanus hirtus Kleine, 1928: 234 (by original (Lycinae) and transferred to the Metriorrhynchinae designation). recently by Bocakova (in press). Samanga and Broxylus are very closely related. They have similar shaped geni­ Diagnosis. Body small. Head without rostrum. talia and large, flat apical palpomeres. These taxa also Antennae slender, weakly serrate, flattened (Fig. 37). have quadrate elytral cells with distinct concave bottoms Maxillary apical palpomere pointed (Fig. 16). Pronotum (these are unique in Metriorrhynchinae) and reduced lat­ with short median areola attached to frontal margin by eral carinae on the pronotum. Thanks to the considerably relatively long keel. Frontolateral carinae less apparent dilated elytra Broxylus has a very characteristic general (Fig. 72). Elytra with three primary longitudinal costae. appearance. The last mentioned character is the only dif­ According to structure of basal part of costae, primary ference between Broxylus and Samanga and therefore I costae 1, 2 and 4 present, costa 3 absent, vestiges of it propose Samanga to be a junior synonym of Broxylus. make base of costa 4 stouter. Secondary costae weak, Elytra of Broxylus fenestratus are less dilated and similar irregular or absent. Transverse costae obtuse, weak, to those of Xylobanus. irregular to well developed. Elytra densely pubescent or Broxylus is very close to Xylobanus from which it dif­ with erect bristles. Phallus lanceolate, apical fourth wider, fers in the shape of its terminal palpomeres. We do not

339 internal sac with two spines. Ovipositor moderately slen­ No genera bearing similar characters were found within der, vagina membranous. the range of Metanoeus. Further characters supporting the Distribution. Oriental Region: Ceylon. Two species classi­ relationship of Metanoeus within Metriorrhynchina are fied in this genus. need. Remarks. The shape of genitalia and pronotal carinae Metriorrhynchus Gemminger et Harold, 1869 indicate a close relationship between Prometanoeus and Metriorrhynchus GemmingeretHarold, 1869: 1629. Cautires. They differ in absence and/or presence of fla- Type species. Lycus parallelus Guerin-Meneville in Boisduval, bellate antennae in male. The flabellate antennae evolved 1835: 114; C. O. Waterhouse, 1878: 101 (by subsequent desig­ several times in Lycidae. The monotypic genera Prome­ nation). tanoeus and Tapromenoeus are very closely related and Metriorhynchus Guerin-Meneville, 1838: 72 (ajunior homonym although they are different in general appearance they of Metriorhynchus Meyer, 1830). have a similar shaped pronotum, pubescence on elytra, Type species. Lycus parallelus Guerin-Meneville in Boisduval, and a similar shaped aedeagus. Bocak & Bocakova 1835: 114; C. O. Waterhouse, 1878: 101 (by subsequent desig­ nation). (1989) basedTapromenoeus on the absence of secondary costae but the analysis showed that the reduction of sec­ Invalid designation. Metriorrhynchus cribripennis C. O. Water­ ondary costae happened several times in the Metrior- house, 1879: 52; Bourgeois, 1891: 347 (by subsequent designa­ rhynchinae and is also often encountered in other sub­ tion). families of Lycidae. Prometanoeus does not have any Dilolycus Kleine, 1926a: 186,syn. n. reliable apomorphic character state that separates it from Type species. Dilolycus lamellatus Kleine, 1926a: 186 (by Tapromenoeus. Therefore I consider Tapromenoeus to be original designation). ajunior subjective synonym of Prometanoeus. Flabelloporrostoma Pic, 1923: 35, hors-texte (as subgenus of Porrostoma), syn. n. Metanoeus C. O. Waterhouse, 1879 Type species. Porrostoma mirabilis Pic, 1923: 35 (by mono- Metanoeus C. O. Waterhouse, 1879: 73. typy). Type species. Lycus (gen. 37) conformis C. O. Waterhouse, 1878: 115; Bourgeois, 1891: 347 (bysubsequentdesignation). Diagnosis.Body large, parallel-sided. Head small, with short, stout rostrum or without rostrum, galeae long, Diagnosis.Body small to medium-sized, elytra at most much longer than mandibles. Antennae serrate in both slightly widened posteriorly. Head small, without sexes or flabellate in male and serrate in female (Fig. 36). rostrum, with large eyes even in females. Male antennal Maxillary palpi slender, long, apical palpomere slender, lamellae long, narrowly attached to basal parts of anten- compressed, pointed at apex (as in Fig. 14). Pronotum nomeres (Fig. 40). Maxillary palpi relatively slender, with seven, in middle sometimes reduced, areoles on disc. apical antennomere parallel-sided. Pronotum with seven Four strong elytral primary costae developed, costa 3 areoles, carinae obtuse (Fig. 73). Phallus wide basally sometimes slightly shortened (by one eighth at most), (Fig. 113). Phallobasal membrane extensive, at least slightly weaker in apical half of elytra. Secondary costae partly sclerotized. Internal sac with three pairs of spines. much weaker. Transverse costae dense, regular, a bit Vagina often considerably sclerotized, form of sclerotisa- stronger than secondary costae, elytral cells transverse, tion highly species specific, narrowed to apex (if vagina regular. Phallus stout, internal sac without spines but with rounded, then basal part of spermaduct is robust), a sclerotised structure (Fig. 114) or similar structure unpaired gland attached basally to apically, basal position accompanied by several pairs of short minute rods correlated with sclerotisation of vagina, lateral glands (Bocak, 1998c, Figs 1, 2). Ovipositor small, only partly large to medium sized, attached ventrally, when vagina sclerotized, with short valvifers (Fig. 134), vagina large, sclerotized, laterally if membranous, basal part of glan­ often at least partly sclerotized (Fig. 148; Bocak 1998c, dular ducts more or less sclerotized. Spermaduct usually Figs 6, 7). extremely long, sometimes of medium length (in species with membranous vagina). Female valvifers considerably Distribution. Australian Region and eastern part of Oriental shortened, modified in plates (Fig. 150). Region: Peninsular Malaysia, Thailand, Laos, Sundas, and the Philippines. Metriorrhynchus sericeus C. O. Waterhouse, 1879 Distribution. Oriental Region. The highest diversity found in is reported from India (Kleine, 1933), butMetriorrhynchus was Borneo, Sumatra, and Peninsular Malaysia. Metanoeus is very not found in the recently collected material from India. Highest rare in western part of Oriental Region. One species is common diversity is found in the Papuan Subregion and Australia, but in Mindanao. Altogether over 20 species are classified in fauna of these regions is badly in need of revision at the species Metanoeus. level. Calder (1998) combined all Australian species Metrior­of Remarks. The very obtuse pronotal carinae and ves­ rhynchus with Porrostoma, because of the doubtful status of the name Metriorhynchus Guerin-Meneville, 1838. tigial valvifers are characteristic of Metanoeus. Similar obtuse carinae are found in some Metriorrhynchus, which Remarks. There are two types of vagina in Metrior­ differ in the much shorter lamellae of male antennae, long rhynchus: M. parallelus (type-species) has a completely slender phallus and only shortened valvifers. The scleroti­ sclerotized vagina but species in the M. thoracicus Fabri- sation of vagina, long processes on scutellum and long, cius, 1801 group have a completely membranous vagina. basally attached male antennal lamellae reminiscent of These groups have a similar shaped paraproctus, very some Papuan representatives of the Metriorrhynchinae.

340 slender, reduced valvifers and a very strong membrane in very short rostrum. Antennae compressed, antennomeres which the ovipositor is enclosed. 3-10 parallel-sided to slightly serrate, never flabellate. Dilolycus was proposed for a species with considerably Pronotum slender, long, apparently narrowed frontally reduced carinae in the middle of the pronotum (Fig. 85). (Fig. 75), only very slender longitudinal areola in middle It has similar shaped male genitalia to the Papuan group of pronotum, sometimes less apparent lateral carinae pre­ of species (as in Bocak, 1998c, Figs 1, 2). Therefore I sent on middle of lateral pronotal margin. Elytra slender, consider Dilolycus to be a junior subjective synonym of with considerably shortened primary costa 1 and with Metriorrhynchus. entire secondary costae. Legs very long, slender, strongly The holotype of Metriorrhynchus mirabilis has dam­ compressed. Male genitalia very diverse in shape, often aged antennae, only the three basal antennomeres of each slender, tube-like, internal sac without any basal spines, antenna remain preserved, antennomere 3 is serrate. Two rarely with strengthened sclerotized parts. Phallobase fragments of antennae were glued on the separate label: often with large, well sclerotized membrane. Ovipositor the first fragment was serrate, corresponding to the with short valvifers, vagina short, accessory glands remaining antennomeres on the holotype, the second frag­ attached laterally, unpaired vaginal gland large, sper- ment flabellate. It must be part of a different specimen, matheca bulb-like. because it is made up by antennomeres 3-11. The name Distribution. Australian Region: Lesser Sundas, Moluccas, Flabelloporrostoma and its generic status are based on and Sulawesi; Oriental Region: Sunda Islands, the Philippines, this incorrectly identified body part. Therefore, I propose Malaysia, Thailand and Indochina. Altogether almost 60 species Flabelloporrostoma to be a junior subjective synonym of are classified in Leptotrichalus with highest diversity in the Metriorrhynchus. Philippines. Lobatang Bocak, 1998 Remarks. The relationship between Leptotrichalus and Lobatang Bocak, 1998b: 190. Lobatang is supported by prolonged galees, very wide Type species. Lobatang papuensis Bocak, 1998b: 190 (by base of male sternum A8, strong, shortened valvifers and original designation). generally short and wide ovipositor and the slender and asymmetrical shape of basal part of phallus. They differ Diagnosis.Body slender, externally similar to Tri- in the prolonged pronotum in Leptotrichalus and the chalus (general body form, shortened first elytral primary sclerotized internal sac in Lobatang. Additionally, the costa, only one longitudinal areola in middle of apical maxillary palpomere is more slender in Leptotri- pronotum, Fig. 9), medium-sized to moderately large. chalus. Head without rostrum. Antennae long, strongly com­ pressed, antennomeres 3-10 almost parallel-sided to ser­ Stadenus C. O. Waterhouse, 1879 rate in both sexes. Maxillary palpi with slender apical pal- Stadenus C. O. Waterhouse, 1879: 61. pomere. Pronotum with one longitudinal, median areola, Type species. Porrostoma dichroum C. O. Waterhouse, 1877: lateral margins parallel-sided or pronotum much wider at 86 (by original designation). base. Elytra flat, with shortened primary costa 1, usually Diagnosis.Body medium-sized. Head small, without with apparent secondary costae, in one species secondary rostrum. Antennae stout, serrate. Maxillary palpi with costae distinctly weaker than transverse ones, irregular, almost parallel-sided apical palpomere, without papillae, interrupted. Legs long, slender, hind trochanters much apex obliquely cut. Pronotum wide, almost straight at wider apically. Phallus robust only at base, slender and basal margin, widest before base, mostly only median apical two thirds very gradually narrowed, internal sac areola present (see remarks), but at least small vestiges of completely sclerotized. Phallobase robust, membrane frontolateral carinae are regularly present (Fig. 76). Pri­ partly sclerotized (Fig. 116). Ovipositor short and wide, mary costae strong, almost completely developed, costa 2 valvifers robust, vagina wide, accessory glands inserted and 4 apparently stronger in apical part, other primary laterally, glandular ducts very short, unpaired apical gland costae very slightly shortened, weaker. Phallus simple, as long as vagina. slender, internal sac without spines or sclerotisation (Figs Distribution. Australian Region: New Guinea, Moluccas, 117, 118). Female genitalia with very robust and wide Sulawesi; Oriental Region: the Philippines. valvifers, which are firmly fused with coxites, vagina Remarks. Leptotrichalus and Lobatang has same membranous, large (Figs 156, 159). shaped female genitalia, especially the shortened, wid­ Distribution. Australian Region. Five species are placed in ened valvifers. Lobatang is characterised by the unique Stadenus. The Afrotropical species previously classified in Sta­ modification of the internal sac, which has not been found denus are now placed in the tribe Slipinskiini (Erotinae) (Bocak in other Lycidae. & Bocakova, 1992). Leptotrichalus Kleine, 1925 Remarks. The very similar shape of male and female genitalia supports the affinity of Stadenus and Porros­ Leptotrichalus Kleine, 1925a: 296. Type species. Metriorrhynchus cyaniventris Kirsch, 1875 (by toma. Porrostoma differs in presence of rostrum, Sta­ original designation). denus has completely fused coxites and valvifers. Stadenus was originally defined by three areoles on the Diagnosis. Body medium-sized to large, slender, pronotal disc, but this character is unreliable. I have slightly widened posteriorly. Head small, without or with studied male and female Stadenus triareolatus in copula

341 (BMNH). They are externally very similar and without 120) . Ovipositor with slender coxites and stout valvifers any doubt they are conspecific. The female specimen has (Fig. 136), vaginamembranous, short. three areoles, the male seven areoles. But even the female Distribution. Australian Region: New Guinea, only four spe­ specimen has some vestiges of frontolateral carinae. cies classified in Metriorrhynchoides. Porrostoma Castelnau, 1838 Remarks. Metriorrhynchoides is closely related to Porrostoma Castelnau, 1838: 26. Porrostoma. Its pronotal carinae are very similar to those Type species.Lycus rufiipennis Fabricius, 1801 (by monotypy). of Porrostoma and additionally they both have a long ros­ Invalid designation. Porrostoma erythropterum Erichson, 1842; trum. Metriorrhynchoides is characterised by a widened C. O. Waterhouse, 1879: 44. apical part of elytra and costae 2 and 3 fused before apex Diagnosis.Body medium sized to large. Head with of elytra. Additionally, Metriorrhynchoides species are a long rostrum (Fig. 10). Antennae serrate to flabellate in bright metallic blue with bright yellow body parts unlike male (Figs 42, 49), serrate in female. Maxillary palpi Australian Porrostoma. All males of Metriorrhynchoides slender, apical palpomere slender, almost parallel-sided, also have serrate antennae. I have not found an apomor- hypopharynx very long, mandibles minute. Pronotum phic feature defining Porrostoma, therefore it is very with seven strongly marked areoles, carinae usually very probable thatMetriorrhynchoides is a subgroup of Por­ strong (Fig. 77). Elytra with entire primary and secondary rostoma and will be synonymised with it in the future. longitudinal costae. Transverse costae regular, dense, Cladophorus Guerin-Meneville, 1830 cells usually strongly transverse (Fig. 52). Male genitalia Cladophorus Guerin-Meneville, 1830: Plate II, Fig. 9. short, robust to slender and long, but regularly simple, Type species. Cladophorus fiormosus Guerin-Meneville, 1830; parallel-sided, internal sac membranous, without spines C. O. Waterhouse, 1878 (by subsequent designation). (Fig. 119). Female genitalia with robust valvifers, very closely attached to coxites, vagina membranous, rela­ Odontocerus Guerin-Meneville, 1830: 72; a homonym of Odon- tocerus Stephens, 1829; C. O. Waterhouse, 1878: 96. tively short (Bocak, 1998c, Fig. 14). Type species. Cladophorus fiormosus Guerin-Meneville, 1830; Distribution. Australian Region. Porrostoma is the com­ Bourgeois, 1892: 495 (by subsequent designation). monest lycid in Australia, where the highest diversity is found. Spacekia Strand, 1936; Bocak, 1998c: 247. Several species are known from New Guinea. Type species. Cladophorus fiormosus Guerin-Meneville, 1830 Remarks. Porrostoma was for long time considered a (Spacekia proposed by Strand, 1936 as a nomen novum forCla- junior synonym of Metriorrhynchus. The name was used dophorus). by C. O. Waterhouse (1877) and later sometimes incon­ Diagnosis.Body medium sized to large, often brightly sistently by Pic. Kleine (1933) listed Porrostoma as a coloured. Head without rostrum. Antennae with very long junior synonym of Metriorrhynchus. Calder (1998) used lamellae in male (Figs 44, 51), serrate in female. Maxil­ Porrostoma as ajunior replacement name for Metriorhyn- lary palpi stout, apical palpomere securiform. Pronotum chus Guerin-Meneville and proposed 85 new combina­ often apparently transverse, with strongly marked seven tions with Porrostoma (only Australian fauna). Bocak areoles (Fig. 79). Elytra with entire primary and secon­ (1998c) replacedMetriorhynchus Guerin-Meneville by dary longitudinal costae. Phallus gradually narrowed to Metriorrhynchus Gemminger et Harold, used Porrostoma apex, internal sac partly sclerotized. Phallobasal mem­ as a valid taxon and differentiated between Metriorrhyn­ brane apparently pigmented and additionally margin of chus and Porrostoma. The identity of most species classi­ phallobase partly sclerotized, with attached membrane. fied in Porrostoma is unknown and this genus is badly in Membrane has a characteristic structure (Fig. 121) and a need of revision. usually reaches halfway along phallus. Ovipositor moder­ Metriorrhynchoides Kleine, 1926 ately wide, simple, valvifers slender (Fig. 141). Vagina Metriorrhynchoides Kleine, 1926a: 118. membranous with short spermaduct and gradually wid­ Type species. Metriorrhynchoides helleri Kleine, 1926a: 119 ened spermatheca (Fig.144) (by original designation). Distribution. Australian Region: New Guinea, Australia. All Diagnosis.Body large, conspicuously widened posteri­ records outside this region refer to other genera. orly. Head small with long, laterally compressed rostrum Remarks. Cladophorus is defined by the form of its (as in Fig. 10). Antennae slender, flattened, serrate in phallus and unique shape of phallobasal membrane (Fig. both sexes (Fig. 43). Apical maxillary palpomere flat, 121) , which has not yet been found in other genera of narrowed to apex. Maxillae long. Pronotum with seven Metriorrhynchinae. At least several dozens of species strongly marked pronotal areoles, median areola large, with this type of aedeagus are known from New Guinea widest in frontal part (Fig. 78). Frontolateral carinae (Kleine, 1926a). On the other hand, most of almost 200 short. Pronotum constricted in basal third. Elytra wide, species previously placed in Cladophorus are not conge­ much wider than abdomen, with elevated humeri. Four neric with the New Guinean species (all species from the primary longitudinal costae strong over whole length, Oriental, Palaearctic and Afrotropical Regions). Kleine costae 2 and 3 sometimes fused before apex. Secondary classified as Cladophorus all species that have slender costae weak but continuous (Fig. 55). Transverse costae antennal lamellae (mostly Cautires). Pic often used the very dense, regular. Elytral cells transverse (Fig. 55). name Odontocerus Guerin-Meneville nec Stephens and Male genitalia slender, internal sac membranous (Fig. these species were transferred to Cladophorus by Kleine

342 (1933). Pic's concept is unclear and his descriptions do allel sided, with four primary and five secondary costae not allow identification bellow subfamily level. extending the whole length, primary costae stronger at Procautires Kleine, 1925 humeri, weak in apical fifth. Transverse costae forming quadrate, rounded cells to slightly longitudinal rounded Procautires Kleine, 1925b: 32 cells. Phallus stout, two plates at base of internal sac, pri­ Type species.Procautires toxopei Kleine, 1925b: 32. mary gonoporus circular, sclerotized (Figs 122, 123). Diagnosis.Body medium sized, moderately robust, Vagina very slender, its wall partly stiffened (Fig. 160). parallel-sided. Head without rostrum. Antennae flabellate Distribution. Australia Region: New Guinea, Moluccas. At in male, serrate in female. Maxillary palpi short, apical present only C. reticulatus is classified here, but several unde­ palpomere securiform, maxillae short, wide, mandibles scribed species exist in collections. very slender, curved. Pronotum with strong carinae and seven distinct areoles. Elytra with four strong longitudinal Remarks. Ditua is similar to Cautiromimus but it dif­ primary costae, secondary costae present on basal fifth fers in serrate male antennae and in the pattern of pro­ and apical two fifths of elytra. Elytral cells minute, notal carinae. An alpha taxonomic revision of material in rounded to square in basal and apical parts of elytra, collections will clarify of their relationship. strongly transverse in the middle. Male genitalia rela­ Ditua C. O. Waterhouse, 1879 tively slender, slightly widened apically, internal sac with Ditua C. O. Waterhouse, 1879: 33. sclerotized basal part and primary gonoporus, phallobasal Type species. Ditua deplanata C. O. Waterhouse, 1878: 100, membrane pigmented, structured (Fig. 126). Ovipositor 109 (by monotypy). slender, coxites almost as long as valvifers, between cox­ fnvalid designation. Ditua dichroma (Boisduval, 1835: 123); ites triangular sclerite, valvifers slender. Vagina short, Bourgeois, 1891: 345. small, membranous, reaching two thirds of length of val- Diagnosis.Body medium sized, parallel-sided. Head vifers. without rostrum. Antennae serrate in both sexes, antenno- Distribution. Australia Region: Australia, New Guinea and meres more acutely triangular in male. Maxillary palpi Moluccas. There are several Afrotropical and Oriental species slender, apical palpomere parallel-sided, slender. Mandi­ now classified in Procautires but they are not congeneric with bles long, slender, curved. Pronotum transverse, with Procautires toxopei and should be classified with Cautires or another closely related genus. These species have less con­ slightly convex frontal margin. Seven areoles on disc, all spicuous secondary costae, but do not share other characters carinae very strong (Fig. 80). Elytra with four entire pri­ with the type species. mary and five secondary longitudinal costae, transverse costae dense, reticulate cell minute, quadrate to rounded. Remarks. Procautires is characterised by a unique Male genitalia with robust phallus, internal sac with basal modification of elytral secondary costae. With Kassemia sclerotized structures and sclerotized primary gonoporus and Cautiromimus it shares the same shape of phallus and (Fig. 125). Ovipositor slender, vaginamembranous. sclerotisation of internal sac and with Cladophorus pig­ mented phallobasal membrane, although it is in this case Distribution. Australian Region: New Guinea. Two species less extensive and does not have the characteristic struc­ are classified here but several undescribed species exist in col­ lections. ture as in Fig. 121. Additionally, f have found a relatively large sclerite between coxites. Very small structure in Remarks. Ditua is reminiscent of Cautiromimus but similar position is present also in some Cautiromimus differs in that male antennae are serrate and in the shape (Fig. 160). The sclerite between bases of coxites is of pronotum. Ditua deplanata is a member of a very reminicent of sclerite found in some species of Xylobanus common lycid Mullerian mimicry complex and f have (Fig. 154), but unlike in Xylobanus, the sclerite is not found in collections several not very closely related spe­ connected with valvifers and is located between bases of cies identified as Ditua dichroma. The name D. deplanata coxites. Therefore f do not consider these structures to be was up to now used as the junior synonym of Ditua homologous. dichroma along with two other names. The status of the synonyms of Ditua dichroma was not checked. Cautiromimus Pic, 1926 Cautiromimus Pic, 1926c: 454 Kassemia Bocak, 1998 Type species. Cladophorus reticulatus Kleine, 1926a: 143 (by Kassemia Bocak, 1998b: 195. monotypy). Type species.Kassemia oculata Bocak, 1998. Diagnosis.Body medium sized, relatively slender. Diagnosis.Body small to medium sized, parallel-sided, Head small, without rostrum. Male antennae flabellate, all representatives dark brown to brown with humeral female antennae serrate. Maxillary palpi with almost par­ quarter to half of elytra light yellow to brown, always dis­ allel sided apical palpomere, without papillae. Middle tinctly much lighter than the rest of elytra. Head small, pronotal areola very narrow, deep, attached to the basal without rostrum, mandibles slender, long, three times margin by very short keel, reaching almost 3/4 of pro­ longer than labrum, apical maxillary palpomere robust, notal length, frontolateral carinae very strong, but weak in securiform, broad. Antennae strongly compressed, flabel­ frontal punctured area, absent in frontal quarter, lateral late in males and serrate in females. Pronotum with seven carinae are not connected with median areola (Fig. 81). areoles. Elytral secondary costae completely absent, pri­ Posterior half of pronotum shining, pubescent. Elytra par­ mary costa 1 shortened, costa 2 and 3 sometimes fused

343 before apex. Legs relatively strong and short, strongly thrix is considered to be a junior subjective synonym of compressed. Male genitalia with relatively short and Pseudodontocerus. robust phallus, apical part widely open, internal sac Oriomum Bocak, 1999 strengthened at base with spine-like structure. Phallobasal Oriomum Bocak, 1999a: 111. membrane never sclerotized. Ovipositor with slender, Type species. Oriomumfemoralis Bocak, 1999a: 111. long valvifers, vagina as long as valvifers, widest at apex, accessory glands inserted laterally. Diagnosis.Body medium-sized, robust. Head rostrate, rostrum reaching to middle coxae. Antennomeres 1 to 5 Distribution. New Guinea, four species known at present (Bocak, 1998b). slender, antennomere 6 of similar shape but slightly ser­ rate, antennomeres 7-9 short, triangular. Bottom side of Remarks. The combination of seven areoles on the antennomeres 1 and 3-4 with very long, dense hairs. Pro­ pronotum, complete absence of elytral secondary costae, notum with 7 areoles. Elytra with four primary costae shape of male genitalia, and shortened primary costa 1 extending whole length, costa 1 weaker in posterior half. confirm the sure identification of Kassemia. The shape of Secondary costae distinct in humeral area, weaker on rest male antennae and shape of male and female genitalia of elytra, cells irregular. Legs significantly modified for show little assotiation with shortened primary costa 1 in clasping of female, with long dense pubesce. Femora of other genera. The shape of phallus indicates a close rela­ fore legs and mid legs with spines basally. Hind legs with tionship with genera of the Cladophorus - Ditua - Cauti- bigger, triangular trochanters. Position of hind process of romimus clade. trochanter corresponds with the position of femoral Pseudodontocerus Pic, 1921 spines on fore and mid legs. Phallus slender, parallel­ Pseudodontocerus Pic 1921b: 12, hors texte. sided, with exposed, dense pubescence on apical part of Type species. Pseudodontocerus pulcher Pic, 1921b: 12 (by internal sac, basal part of internal sac without sclerotisa- monotypy). tion. Female unknown. Carathrix Kleine, 1926a: 149, syn. n. Distribution.Australian Region: New Guinea. Monotypic. Type species. Carathrix pilosus Kleine, 1926a: 149 (by original Remarks. Oriomum differs from the remaining Metri- designation). orrhynchinae genera in the unique shape of its elytra with Diagnosis.Body slender, medium sized. Head without entire but apically distinctly weaker first primary costa, rostrum. Lamellae of male antennae extremely long (Fig. the presence of spines on hind margins of fore and mid 45). Female antennae flabellate, very slender, antenno- femora, hind legs with acutely projected triangular tro­ meres 2 and 3 with lamella shorter than antennal trunk chanters and opistognathous head with extremely long (Figs 46, 47), lamellae gradually prolonged, and on rostrum reaching the coxae of the mid legs, and with middle and apical antennomeres clearly longer than trunk. antennomeres of a very unusual shape. Oriomum seems to Maxillary palpi long, with slender palpomere 2 and very be closely related to the Porrostoma clade, but for the short palpomeres 1 and 3, apical palpomere broad, with final decision on this a female is necessary. papillae on apical edge (Fig. 18). Labial palpi tiny, with Cladophorinus Kleine, 1926 broad apical palpomere and papillae on its margin. Pro­ notum with seven areoles (Fig. 83), carinae in middle of Cladophorinus Kleine, 1926a: 149. Type species. Cladophorinus cyaneus Kleine, 1926a: 150 (by disc sharp and apparent, more obtuse on margins. Elytra original designation). parallel-sided, slightly widened at apex. Four primary and five secondary costae strong over whole elytral length, Diagnosis.Body medium sized, slender, slightly wid­ transverse costae dense, slightly irregular, elytral cells ened in posterior quarter, whole body a shiny metallic mostly quadrate, often rounded (Figs 52, 54). Legs blue. Head without rostrum. Male antennae flabellate, slender (Fig. 161). Phallus relatively robust, with basal lamellae slightly longer than trunk of antennomere, spines and extensively sclerotized internal sac (Figs lamellae originate from apical part of antennomere. Max­ 127-128). Ovipositor very slender (Fig. 139), vagina con­ illary palpi moderately long, apical palpomere securiform. stricted, basal part membranous, apical two thirds partly Mandibles curved, sickle-like, long. Pronotum with seven sclerotized (Fig. 143). areoles, median areola wide, carinae obtuse, partly less distinct. Scutellum deeply emarginate at apex. Elytra with Distribution.Australian Region: New Guinea. four entire longitudinal primary and five secondary Remarks. The flabellate female antennae are the auta- costae, primary costa 1 weaker. Secondary costae consid­ pomorphy of Pseudodontocerus and this character is not erably weaker, transverse costae obtuse, irregular. Male present in other Lycidae. Pseudodontocerus is character­ genitalia slender, phallus widened apically, internal sac ised by the extremely long antennal lamellae in males, partly sclerotized (Fig. 124). Female unknown. weaker lateral and frontolateral pronotal costae, and a Distribution.Australian Region: New Guinea. Monotypic. slender body shape. I examined type species of Carathix and Pseudodontocerus as well as several closely related Remarks. Only the damaged holotype of Cladophor­ undescribed species, and have not found any difference inus cyanescens was available for this study. Therefore supporting their separate generic status. Therefore, Cara­ the genus was not coded into the matrix and is now classi­ fied in Metriorrhynchina on the basis of the shape of male genitalia and general appearance. The species is easily

344 recognisable by its metallic shine, seven pronotal areoles distinct antennal tubercles. Eyes small, hemispherically and flabellate antennae. prominent. Antennae very slender, compressed (Fig. 48). Mimoxylobanus Pic, 1921 Palpi slender, apical maxillary palpomere pointed. Pro­ notum with considerably projected posterior angles. Pro­ Mimoxylobanus Pic, 1921b: 11, hors texte. notal carinae absent and only an indistinct median areola Type species. Mimoxylobanus angustatus Pic, 1921b: 11 (by monotypy). (Fig. 84). Elytra narrow, 5.5 times longer than their com­ bined width at humeri. Each elytron with four, slender Diagnosis.Body small, slender, almost parallel-sided. primary costae, costae weak and hardly distinguishable in Head without rostrum. Male antennae apparently com­ apical third. Secondary costae absent, transverse costae pressed, antennomeres 3-10 parallel-sided. Maxillary sparse, elytral cells mostly longitudinal (Fig. 56). Phallus palpi short, robust, apical palpomere weakly securiform, short, stout, with simple sclerotisation of internal sac (Fig. short. Pronotum only slightly wider at base than long at 129). Female unknown. midline, lateral margins almost parallel-sided, straight. Seven areoles on disc of pronotum, lateral carinae obtuse Distribution. Australian Region: New Guinea. Monotypic genus. (Fig. 86). Elytra with four primary longitudinal costae, Remarks. Only the holotype of Malacolycus paululus secondary costae absent, transverse costae irregular, is known. The pronotal carinae are considerably reduced reticulate cells mostly quadrate. All costae distinct, but and simple male genitalia are very similar to those of low in profile. Male genitalia slender, lanceolate, with Xylobanomimus. Therefore, Malacolycus is placed in the protracted apical part of internal sac (Figs 131, 132). Metriorrhynchina. The body form and structure of elytra Female unknown. resembles that of Xylobanomorphus, but they differ in Distribution.Australian Region: Sumbawa. Monotypic. male genitalia (Figs 129, 130). Remarks. Only the holotype of Mimoxylobanus angus­Xylobanomorphus Kleine, 1935 tatus was available. In general appearance and shape of Xylobanomorphus Kleine, 1935: 316. phallus this species resemblesXylobanus. The exact posi­ Type species.Xylobanomorphus transformis Kleine, 1935: 316 tion of this genus within the Metriorrhynchina is because (by monotypy). it differs substantially in the structure of internal sac. Diagnosis.Body small, very slender, delicate, covered Xylobanomimus Kleine, 1926 with erect setae. Head small, without rostrum, eyes very Xylobanomimus Kleine 1926a: 166. small, their interocular distance 3.2 times longer than Type species. Xylobanomimus papuensis Kleine, 1926a: 166 maximum diameter. Antennomeres 3-10 slender, parallel­ [by original designation, mentioned as X. papuanus (sic!)] sided. Maxillary palpi slender, apical palpomere pointed. Diagnosis.Body small, slender, delicate. Head without Pronotum with almost parallel-sided lateral margins and rostrum. Male antennae flabellate, lamellae slender, up to projected posterior angles. Only median areola distinct, four times longer than trunk of antennomere, attached to lateral carinae obtuse, frontolateral carinae absent. Elytra base on basal segment or middle part on more distal very slender, with four primary longitudinal costae, sec­ antennomeres. Maxillary palpi slender, long, apical pal­ ondary costae absent. Transverse costae scarce, elytral pomere parallel-sided, apex pointed. Pronotum with cells longitudinal (Fig. 56). Male genitalia slender, seven areoles, middle areola large, lateral carinae obtuse. phallus lanceolate, internal sac extensive with ring­ Elytra with four primary costae, vestiges of secondary shaped sclerotized structure (Fig. 130). Female unknown. costae at humeri. Elytral areoles quadrate to longitudinal. Distribution.Australian Region: New Guinea. Elytra with sparse, long, erect setae. Phallus robust, Remarks. At present, only the holotype of Xylobano­ parallel-sided, with simple sclerotisation of internal sac morphus transformis is known. This species resembles (as in Fig. 129). Female unknown. Xylobanomimus and Malacolycus in general appearance, Distribution. Australian Region: New Guinea. Monotypic but differs in the sclerotisation of the internal sac. Consid­ genus. ering its similarity to Xylobanomimus, the absent fronto­ Remarks. Xylobanomimus has very similar male geni­ lateral pronotal carinae can be hypothesised as second­ talia, general appearance and structure of elytral cells to arily reduced. Several common characters and similarity Malacolycus, but differs in the flabellate male antennae in general appearance support the affinity of and strong fTontolateral carinae on the pronotum. Malaco­ Malacolycus, Xylobanomimus and Xylobanomorphus. All lycus can be hypothesised as a sister group of Xyloba­ these taxa are known from single specimens of males. nomimus. Xylobanomimus is classified with Metriorr- Further material is needed for study and provisionally hynchina. Xylobanomorphus is classified incertae sedis in Metrior- rhynchina. Malacolycus Kleine, 1943 Malacolycus Kleine, 1943: 151 Taxon dubium Type species. Malacolycus paululus Kleine, 1943: 151 (by Falsolucidota Pic, 1921 monotypy). Falsolucidota Pic, 1921b: 9, hors texte. Diagnosis.Body small, very slender, only slightly Type species. Falsolucidota testaceicollis Pic, 1921b: 9 (by monotypy). sclerotized. Head small, with convex Irons and small, but

345 Remarks. The identity of the monotypic genus Falso- 11. Elytral secondary costae only absent in middle of elytra, lucidota is unknown. The type specimen was not found in present on basal quarter and apical fifth, internal sac with Pic’s collection in Paris and the original description does sclerotized circular gonoporus and basal plates (Fig. 126), elytral cells in middle of elytra transverse . . . Procautires not help in the identification of this taxon. - Elytral secondary costae completely absent or at most vestiges Key to Metriorrhynchinae genera presentonsmallareaatthebaseofpronotum ...... 12 1. Elytral primary costa 1 entire and strong in whole length, 12. Pronotum with small, very slender median areola, connected weaker in apical half or very slightly shortened; always to frontal margin via strong keel, or median areola very reachingatleastbeyondhalfwayalongelytra ...... 10 obtuse to absent (Fig. 67), frontal margin formed by two - Elytral primary costa 1 considerably shortened, never reaching strong carinae (Fig. 67), elytral areoles quadrate, with mat, beyond half way along elytra, typically not longer than one velvet-like bottoms, found only in Madagascar ...... third of elytral length (Fig. 9) ...... 2 ...... Caenioxylobanus 2. Pronotum with seven areoles (the same type of arrangement - Pronotum with different arrangement of carinae (Figs 57, 60, of pronotal carinae as in Figs 69, 71 or 80), only three longi­ 69,70,72,84),frontalmarginofpronotumsimple .... 13 tudinal elytral costae (costa 2, 3 and 4) present in the middle 13. Male genitalia with parameres (as in Fig. 91), at most para- of each elytron; costa 1 shortened (as Fig. 9), antennae fla- meres shortened ...... Xylobanellus bellate (as Fig. 38) ...... Kassemia -Parameresabsent(Figs 92-130) ...... 14 - Pronotum with less than seven areoles (as in figs 62-66) . . 3 14. Pronotum with five areoles, pronotal carinae slender, 3. Pair of vaginal glands inserted dorsally, directed apically or straight (as in Fig. 60), phallus very slender with one com­ laterally (Bocak, 1998b: Figs 68-70), apical part of phallus plex spine at base or in middle part of phallus, apical part narrower than middle part, ventrally open, only dorsal part oftenrotated(Fig.94) ...... Wakarumbia sclerotized(Figs 103-106) 4 - Pronotum with differently arranged pronotal carinae, always - Pair of vaginal glands inserted laterally (Fig. 157), apical part at last vestiges of frontolateral carinae present (Figs 69-83) of phallus as wide or wider than the middle (Figs 100-102, or only one unapparent median areola present (Fig. 84), 115,116) 9 internal sac with two spines in various positions (Figs 4. Pronotum with five areoles (similar to Fig. 70) or with two 107-112,121-128)orwithoutspines(Fig.130) 15 anteriorly divergent longitudinal carinae (Fig. 66), phallus 15. Most elytral cells distinctly longitudinal (Fig. 56), body very withpigmenteddorsalkeel(Figs 105,106) 5 slender, minute, delicate, pronotum costae obtuse, indistinct, - Pronotum with one lanceolate longitudinal areola attached to pronotum with one inconspicuous areola (Fig. 84) or ves­ frontal and basal margins of pronotum at one point (Figs tiges of lateral and frontolateral carinae present ...... 16 62-65) ...... 6 - Elytral cells quadrate to transverse, body medium sized to 5. Pronotum with five areoles (similar to Fig. 70) ...... moderately large, pronotum always at least with vestiges of ...... Schizotrichalus frontolateralcarinae(Figs 69-83) 18 - Pronotum with two anteriorly divergent longitudinal carinae 16. Phallus parallel-sided with rounded apex, internal sac with (Fig.66) Eniclases twosclerotizedrods(Fig.128) ...... 17 6. Male antennae flabellate (similar to Figs 44, 47) ...... - Phallus slender, narrowed to apex, internal sac with ring­ ...... Flabellotrichalus shaped sclerotised structure (Fig. 130) ...... - Male antennae serrate or antennomeres parallel-sided (Figs ...... Xylobanomorphus 34,35) 7 17. Male antennae flabellate, vestiges of pronotal lateral and 7. Vagina with two lateral pockets medially and with an frontolateral costae present...... Xylobanomimus unpaired very slim and long gland basally (Bocak, 1998b), - Male antennomeres parallel-sided (Fig. 48), only indistinct valvifers slender, usually free, sometimes connected in basal median areola present (Fig. 84) ...... Malacolycus part ...... Microtrichalus 18. Bottom of elytral cells shiny, apparently concave, quadrate - Vagina without lateral pockets, unpaired gland absent, val­ (Fig. 90), elytra more or less dilated, convex, pronotal lat­ vifers robust, connected subbasally (Bocak, 1998b, Fig. 64) eral carinae always indistinct to absent (Fig. 70), male geni­ ...... Trichalus talia as in Fig. 111 Broxylus 8. Maxillary and labial palpomeres with distinct sensilae, apical - Bottom of elytral cells mat, pubescent, elytra parallel-sided to palpomeres broad, securiform (similar to Fig. 18), apical very slightly dilated, f la t ...... 19 part of phallus projected ventrally, internal sac very com­ 19. Elytra with three longitudinal primary costae running whole plex, sclerotized(Figs 100-101) Diatrichalus length of elytra ...... Prometanoeuspart - Maxillary and labial palpomeres slender, internal sac mostly - Elytra with four longitudinal primary costae running whole without sclerotisation, sometimes completely sclerotized length of elytra ...... 20 (Figs 115-116) ...... 9 20. Pronotum with seven areoles, lateral carinae regularly well 9. Pronotum very slender, longitudinal areola very slender (Fig. developed, seldom lateral carinae more obtuse, internal sac 75), internal sac membranous or at most with slender sclero- with pair of spines apically, male antennae serrate to flabel­ tizedrods ...... Leptotrichalus late (as in Figs 38, 39) Xylobanus - Pronotum approximately as wide as long, internal sac com­ - Pronotum with seven areoles, lateral carinae obtuse, internal pletely sclerotized, very slender, basally attached to phallus sac without spines, and the internal sac with a very slender (Fig. 116) ...... Lobatang apicalpart(Figs131-132) ...... Mimoxylobanus 10. Elytral secondary costae absent at least medially (Figs 56, 21. Male genitalia with parameres (Fig. 91), unpaired vaginal 87,88,90) 11 gland inconspicuous, pronotum as in Fig. 58 .... Conderis - Elytra with very distinct secondary costae on whole elytra - Male genitalia without parameres (Figs 92-130), unpaired (Figs 44, 45, 48) ...... 21 vaginal gland large (Figs 163-159), pronotum with carinae as in Figs 59-86 ...... 22 22. Elytra with three longitudinal primary costae running whole length of elytra ...... Prometanoeus part

346 - Elytra with four longitudinal primary costae running whole 33. Phallus lanceolate, internal sac with pair of spines apically length of elytra ...... 23 (Fig. 108), no body part shiny metallic blue .... Cautires 23. Frontolateral carinae on pronotum absent (Figs 59-60, 68) . - Phallus parallel-sided, tubular, shorter, body often a shiny ...... 24 metallic blue ...... 34 - Frontolateral carinae on pronotum present (Figs 69-74, 34. Pronotal carinae robust but obtuse, indistinct, male antennae 76-83) ...... 28 with long, relatively slender lamellae (Fig. 40), male geni­ 24. Pronotum with very obtuse frontal angles, distinctly nar­ talia parallel-sided (Fig. 113) or robust basally, internal sac rowed anteriorly, with one small, slender median areola with several pairs of processes (Fig. 113), ovipositor with connected with frontal margin by strong keel, lateral carinae considerably shortened valvifers (Fig. 150), lateral glands absent (Fig. 68), phallus lanceolate, internal sac with pair of sometimes attached dorsally and directed upwards, sper- spines (Fig. 107), male antennae flabellate as in Fig. 38 ... maduct often extremely long ...... Metanoeus ...... Bulenides - Pronotal carinae moderately strong and well marked, some­ - Pronotum with conspicuous frontal angles, lateral carinae pre­ times partly reduced, male genitalia never with apical proc­ sent to vestigial, median areola more extensive (Figs 59-61) esses, usually with variously shaped plates or spines, val­ ...... 25 vifers never considerably shortened (Figs 140, 152, 160), 25. Lateral carinae apparent, median areola rhomboidal (Figs lateral glands attached laterally or lateroventrally, sper- 59-60), pronotal carinae usually straight, slender, phallus maductmoderatelylong ...... 35 slender as in Fig. 92 or robust (Fig. 95, 96), with paired 35. Body often large, slender, internal sac of phallus spirally spines or spines arranged linearly, male genitalia slender, sclerotised (Fig. 114), or with straight sclerotisation and sev­ spermathecabulbous...... 26 eral pairs of adjacent, small, simple spines (Bocak, 1998c, - Lateral carinae vestigial, male genitalia as in Figs 97-99, sper- Figs 1, 2), ovipositor very small, lightly sclerotized, partly matheca tubular, very long ...... Enylus membranous (Fig. 134), vagina often considerably 26. Elytral primary costa 3 reaching mid way along elytra, sclerotized ...... Metriorrhynchus spines on internal sac arranged linearly, bases of valvifers - Body mostly medium-sized, phallus without any slender, rod­ with various processes, male antennae as long as body .... like sclerotized structure (Figs 121-125, 127-128), ovi­ ...... Hemiconderis positor well sclerotized (Figs 140), vagina never strongly - Elytral primary costa 3 reaching beyond middle of elytra, usu­ sclerotized ...... 36 ally complete although weaker apically, spines on internal 36. Membrane of phallobase extensive, pigmented, with charac­ sac as in Figs 92-96, bases of valvifers without any process, teristic structure (Fig. 121), phallus narrowed to apex , male male antennae shorter than body ...... 27 antennae with very long lamellae (Figs44,51) ...... 27. Body very slender, fragile, almost parallel-sided, secondary ...... Cladophorus costae very weak, phallus short, slender (Figs 95-96) .... - Phallobasal membrane very soft, small, without any structure ...... Achras (Figs 122-125), never considerably narrowed at apex, male - Body small, but moderately robust, widened posteriorly, sec­ antennae serrate or with short lamellae, lamellae at most ondary costae well developed, phallus slender (Figs 92-93) twicelengthofbodyofantennomere ...... 37 ...... Synchonnus 37. Body metallic blue, male antennae flabellate, phallus slender 28. Head with long rostrum (Fig. 10), phallus slender, tubular, (Fig. 124) ...... Cladophorinus internal sac completely membranous (Figs 117-120) .. 29 -Bodyotherwisecoloured,nevermetallicblue ...... 38 - Head without rostrum (Fig. 11) or head only slightly pro­ 38. Male antennae serrate, male genitalia as in Fig.125 ...... tracted, internal sac usually at least partly sclerotized (Figs ...... Ditua 107-114,121-128) 31 - Male antennae flabellate, male genitalia as in Figs 122-123 . 29. Elytra considerably widened posteriorly, both ventral and ...... Cautiromimus dorsal surface ofbody a distinctly shiny metallic blue .... Genus excluded from the subfamily ...... Metriorrhynchoides - Elytra slender, parallel-sided to slightly widened posteriorly, Metriorrhynchinae sometimes only ventral surface of body a shiny metallic Lycoprogenthes Pic, 1915 blue ...... 30 Lycoprogentes Pic, 1915: 6 30. Antennae with slender, long basal antennomeres and short, Type species. Lycoprogentes pouilloni Pic, 1915: 6 (by mono- triangular apical antennomeres, antennae with long, dense typy). setae, fore and mid femora with spines, hind trochanters Pseudosynchonnus Pic, 1922 - syn. n. triangular...... Oriomum Pseudosynchonnus Pic, 1922: 13, hors texte. - Antennae serrate to flabellate, all antennomeres subequal, Type species. Pseudosynchonnus testaceus Pic, 1922: 13 without long setae, femora without spines, trochanters (hereby designated). slender...... Porrostoma 31. Male antennae with extremely long and very slender, fragile Parapyropterus Kleine, 1926c: 104- syn. n. lamellae (Fig. 45), female antennae with short lamellae Type species. Parapyropterus nigrostriatus Kleine, 1926c: 104 (Figs 46-47) ...... Pseudodontocerus(by monotypy). - Male antennae serrate to flabellate (Figs 38-43) or with long Protaphes Kleine, 1926b: 363 - syn. n. but moderately slender lamellae (Figs 44, 51), female Type species. Protaphes confertus Kleine, 1926b: 364 (by antennae never flabellate ...... 32 original designation). 32. Phallus tubular, with completely membranous internal sac, valvifers fused with coxites, lateral and frontolateral pro­ Pseudosynchonnus is here transferred from the Metrior­ notal carinae sometimes vestigial ...... Stadenus rhynchinae to Erotinae and Lycoprogenthes from - Phallus different, never with completely membranous internal Calochrominae to Erotinae. I have compared Pseudosyn­ sac, valvifers free, frontolateral and lateral pronotal carinae chonnus, Lycoprogenthes, Parapyropterus and Protaphes always distinct ...... 33 (the last two already sooner classified in Erotinae) and I

347 have not found any substantial difference between them. Schizotrichalus Kleine, 1926a: 2, p. 183.Trichalus nigrescens Therefore, I consider genera Pseudosynchonnus, Parapy- C.O. Waterhouse, 1879: Holotype, male, “Mysool” (BMNH). ropterus, and Protaphes to be junior synonyms of Lyco- Eniclases C. O. Waterhouse, 1879: 66.Lycus (gen. 35) luteolus progenthes. C.O. Waterhouse, 1878: Holotype, male, “Bowring 63.47*” (BMNH). Appendix 1: List of type material studied. = Trichalolus Pic, 1923: 36, hors texte. Trichalus (Trichalolus) Subfamily Metriorrhynchinae Kleine, 1926 apertus Pic, 1923: Holotype, female, “Andai, Nle Guinée” (MNHP). Tribe Conderini Bocak et Bocakova, 1990 Subtribe Metriorrhynchina Kleine, 1926 Conderis C. O. Waterhouse, 1879: 59. Calopteron signicolle Kirsch, 1975: Holotype, 1 spec., “Malacca” (SMTD). Bulenides C.O. Waterhouse, 1879: 34: Lycus (gen. 21) obso- =Pseudoconderis Pic, 1921a: 8, hors texte.Pseudoconderis gor- letus C. O. Waterhouse, 1878: Holotype, female, “Java” hami Pic, 1921a, Holotype, female, “India, coll. Armitage (BMNH). (MHNP). Caenioxylobanus Pic, 1922: 13, hors texte. Caenioxylobanus Xylobanellus Kleine, 1930b: 171. Xylobanellus atricolor ater Pic, 1922: Holotype, female, “Madagascar, collection Le Kleine, 1930b: Holotype, female, “Malay. Penin. Selangor, Moult, Janvier” (MHNP). Bukit Kuta” (BMNH). Cautires C.O. Waterhouse, 1879: 36.Lycus (gen. 22) excellens = Rossioptera Kasantsev, 1988: 169.Eros erythropterus Baudi, C. O. Waterhouse, 1878: Holotype, male, “Sar.” (= Sarawak) 1871. The type ofE. erythropterus was not studied. (BMNH). = Chuzenjianus Nakane, 1969: 176. Conderis tenuis Kono, Prometanoeus Kleine, 1925c: 133. Prometanoeus ochraceus 1932. The type species was not examined. Kleine, 1926b: Holotype, male, “Ceylon, G. Lewis, 1910-320; Dikoya, 3800-4200ft, 6. xii. 81-16. i. 82” (BMNH) Tribe Metriorrhynchini Kleine, 1926 Tapromenoeus Bocak et Bocakova, 1989: 327.Xylobanus hirtus Subtribe Hemiconderina Bocak et Bocakova, 1990 Kleine, 1928: 234. Paratype, male, “Dikoya, Ceylon” (MIZW). Synchonnus C. O. Waterhouse, 1879: 59. Porrostoma clien- Broxylus C.O. Waterhouse, 1879: 20.Calopteron pfeifferi C. O. tulum C. O. Waterhouse, 1877: Syntype, male, “Moreton Bai” Waterhouse, 1878: Holotype, male, “Celebes, Wallace” (BMNH). (BMNH). Strophicus C. O. Waterhouse, 1879: 73. Strophicus nigellus C. = Samanga Pic, 1921b: 9, hors texte.Samanga fenestrata Pic, O. Waterhouse, 1879: Holotype, female, “Mysool” (BMNH). 1921b: Holotype, male, “Samanga, S. Celebes, Nov. 1895, H. Hemiconderis Kleine, 1926a: 162. Hemiconderis explicatus Fruhstorfer” (MHNP). Kleine, 1926a: ?Paratype, male, “Mamberamo R., W. G. v. Xylobanus C. O. Waterhouse, 1879: 38.Lycus costifer Walker, Heuer, 20. xii. -21.i. Pionierbivak” (MIZW). 1858: Holotype, male, “Ceylon” (BMNH). Wakarumbia Bocak, 1999b: 166.Wakarumbia gracilis Bocak, Mimoxylobanus Pic, 1921b: 11, hors texte. Mimoxylobanus 1999: Holotype, male, “Sulawesi SE, Buton Island, angustatus Pic, 1921b: Holotype, male, “Sumbawa, Doherty” Wakarumba, 3.-7. ii. 1994, M. Strba and I. Jenis lgt.” (LMBC). (MHNP). Achras C.O. Waterhouse, 1879: 61.Porrostoma limbatum C. O. Metriorrhynchus Gemminger et Harold, 1869: 1629. The type Waterhouse, 1877: Holotype, male, “Austral. 73-6” (BMNH) of Metriorhynchus parallelus Guérin-Méneville, 1838 was not found in either Brussels or Paris Museums and was very Subtribe Trichalina Kleine, 1928 probably destroyed during a fire in the Brussels Museum (L. Enylus C.O. Waterhouse, 1879: 72. Enylus segregatus C. O. Baert, pers. comm.). Waterhouse, 1879: Holotype, female, “Mysool” (BMNH). = Metriorhynchus Guérin-Méneville, 1838: 72 (unavailable Diatrichalus Kleine, 1926a: 167. Diatrichalus xylobanoides senior objective synonym of Metriorrhynchus Gemminger et Kleine, 1926a: Holotype, male, “D.N. Guinea, 285, Hun- Harold. steinspitze, 3. iii. 43, Kais. Augustafl. Exp., Burgers S.G.” = Flabelloporrostoma Pic, 1923: 35, hors-texte. Porrostoma (NHMB). (Flabelloporrostoma) mirabilis Pic, 1923: Holotype, female, = Mimotrichalus Pic, 1930: 92, hors texte. Mimotrichalus tenim- “N. S. Wales” (MHNP). berensis: Holotype, male, “Tenimber Is, Coll. by W. Doherty” Leptotrichalus Kleine, 1925a: 296. Metriorrhynchus cyaniven- (MNHP). tris Kirsch, 1875: Holotype, female, “Malacca” (SMTD). Trichalus C. O. Waterhouse, 1877: 82.Trichalus flavopictus C. Lobatang Bocak, 1998b: 190.Lobatang papuensis Bocak, O. Waterhouse, 1877: Holotype, male, “Port Bowen, 75.22” 1998b: Holotype, male, “Irian Jaya: Jayawijaya, Bime, (BMNH). 1600-1900 m,11.9. 1993, leg. A. Riedel” (SMNS). = Xantheros Fairmaire, 1877: 167. The type of Xantheros Metanoeus C.O. Waterhouse, 1879: 73. Lycus (gen. 37) con- ochreatus Fairmaire, 1877 was destroyed during the World War formis C.O. Waterhouse, 1878: Holotype, male, “Borneo” II in Hamburg (Bocak, 1998b). (BMNH). Microtrichalus Pic, 1921b: 9, hors texte.Microtrichalus singu- Stadenus C.O. Waterhouse, 1879: 61.Porrostoma dichroum C. laris Pic, 1921b: Holotype, male, “?Java” (MHNP). O. Waterhouse, 1977: Holotype, female, “K. George’s Pound, = Falsoenylus Pic, 1926a: 29, hors texte. Falsoenylus 75-36” (BMNH). basipennis Pic, 1926a: Holotype, female, “Baguio, Luzon” Porrostoma Castelnau, 1838: 26.Porrostoma rufipennis Fabri- (MNHP). cius, 1801: The specimen compared with Fabricius’ type was Flabellotrichalus Pic, 1921b: 9, hors texte. Flabellotrichalus available for study (O. Martin, pers. comm.). notatithorax Pic, 1921b: Syntype, male, “Batjan, Aug-Septbr.” Metriorrhynchoides Kleine, 1926a: 118.Metriorrhynchoides (MHNP). helleri Kleine, 1926a: Paratype, male, “K. Wilhelmland, = Villosotrichalus Pic, 1921b: 9, hors texte. Villosotrichalus Bongu” (MIZW). reductus Pic, 1921b: Syntypes, male, female, “Andai, Nlle Gui­ Oriomum Bocak, 1999a: 111. Oriomum femoralis Bocak, nee” (MHNP). 1999c: Holotype, male, “New Guinea: Papua, W. District Govt. Sta. 26.-28. x. 1960, J. L. Gressitt Collector” (BPBM).

348 Cladophorus Guerin-Meneville, 1830: Plate II, Fig. 9. The type REFERENCES of Cladophorus formosus Guerin-Meneville, 1830 was not Bocak L. 1998a: New and little known species of Conderini and found in the Brussels or Paris Museums and was very probably Ateliini (Coleóptera: Lycidae). Acta Univ. Pal. Olom., Bio­ destroyed during a fire in the Brussels Museum (L. Baert, pers. lo g ica l: 17-26. comm.). Bocak L. 1998b: A generic revision and phylogenetic analysis = Odontocerus Guerin-Meneville, 1838: 72 (a homonym of of the subtribe Trichalinina (Coleoptera: Lycidae: Metrior- Odontocerus Stephens, 1829 (Trichoptera)). rhynchini). ActaSoc. Zool. Bohem. 62: 167-200. =Spacekia Strand, 1936: 169; Bocak, 1998c: 247 (a objective Bocak L. 1998c: Nomenclatural notes on taxa of the family synonym of Metriorrhynchus Guerin-Meneville, 1830). Lycidae described by Guérin-Méneville (Insecta: Coleoptera). Cladophorinus Kleine, 1926a: 149. Cladophorinus cyanescens Ann. Zool. 48: 245-251. 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Bocak L. 1999b: New taxa of the subtribe Hemiconderinina Kassemia Bocak, 1998b: 195.Kassemia oculata Bocak, 1998b: (Coleoptera: Lycidae) from Indonesia and New Guinea. Ento­ Holotype, male, “New Guinea, NE Wau, 1200-1300 m, 22. mol. Bl. 95: 166-170. x.1965, J. Sedlacek Collector Bishop Mus.” (BPBM). Bocak L. 2000a: A revision of the genus Diatrichalus Kleine Pseudodontocerus Pic, 1921b: 12, hors texte. Pseudodontocerus from the Philippines. Raffl. Bull. Zool. 48: 11-16. pulcher Pic, 1921b: Holotype, female, “Humboldt Bay, N. Bocak L. 2000b: To the knowledge of the genus Leptotrichalus Guinea, Doherty” (MHNP). Kleine from Sumatra, Borneo and continental Asia (Coleop­ = Carathrix Kleine, 1926a: 149. Carathrix pilosus Kleine, tera: Lycidae). Ann. Soc. Entomol. Fr.(N. S.) 36: 171-184. 1926a: Holotype, male, Paumonu Riv. ix. - xii. 92” (MSNG). Bocak L. 2000c: A revision of the genus Wakarumbia (Coleop­ Xylobanomimus Kleine, 1926a: 166.Xylobanomimus papuensis tera: Lycidae). Eur. J. Entomol. 97: 271-278. Kleine, 1926a, Holotype, male, “N. Guinea, Dilo, Loria, vi-vii Bocak L. 2000d: A revision of the Microtrichalus from 90” (MSNG). Sumatra, with notes on Oriental and Australian species (Cole­ Xylobanomorphus Kleine, 1935: 316. Xylobanomorphus trans- optera: Lycidae). ActaSoc. Zool. Bohem. 64: 3-16. formis Kleine, 1935: Holotype, male, “Papua: Mt. Tafa, 8500ft, Bocak L. 2001a: A revision of the genus Diatrichalus Kleine ii. 1934, L. E. Cheesman, B. M. 1934-244” (BMNH). from New Guinea (Coleoptera: Lycidae). Stuttgarter Beitr. Malacolycus Kleine, 1943: 151. Malacolycus paululus Kleine, Naturk., Ser. A. 622: 1-32. 1943: Holotype, male, “N. Guinea, Biro, 901, Friedrich-Wilh.-hafen” (HNHM). Bocak L. 2001b: New species of the genus Wakarumbia from Dilolycus Kleine, 1926a: 186.Dilolycus lamellatus Kleine, Sulawesi (Coleoptera: Lycidae). RafflesBull. Zool. 49:1-9 1926a: Holotype, male, “N. Guinea, Dilo, Loria, vi-vii 90” Bocak L. & Jass R. (in press): Revision of the genus Broxylus Waterhouse (Coleoptera, Lycidae). Deutsche Entomol. (MSNG). Zeitschr. Genera classified outside the Metriorrhynchinae Bocak L. & Bocakova M.1987a: Notes on the taxonomy of Lycoprogentes Pic, 1915: 6. Lycoprogenthes Pic, 1915: 6. Lyco- some European species of the family Lycidae (Coleoptera). progenthes pouilloni Pic, 1915: Holotype, female, “Darjeling, ActaEntomol. Bohemoslov. 84: 111-121. Pouillon” (MNHP). Bocak L. & Bocakova M. 1987b: A review of Lycidae (Cole­ = Pseudosynchonnus Pic, 1922: 13. Pseudosynchonnus testa- optera) from Taiwan, with a description of a new species. ceus Pic, 1922: 13: Holotype, male, ”Perak” (MHNP). ActaEntomol. Bohemoslov. 84: 273-286. = Protaphes Kleine, 1926b: 363.Protaphes confertus Kleine, Bocak L. & Bocakova M. 1989: Contribution a l’étude des 1926b: 364: Holotype, male, “Mindanao, Surigao” (MIZW). Lycides de Pile Sri Lanka, III. Cladophorini (Coleoptera, Lycidae). DeutscheEntomol. Zeitschr. 36: 321-328. ACKNOWLEDGEMENTS. This study was made possible by a Bocak L. & Bocakova M. 1990a: Revision of the supergeneric grant from the Czech Grant Agency (Project 206/98/0539). The classification of the family Lycidae (Coleoptera). Polskie following individuals and institutions are gratefully acknowl­ Pismo Entomol. 59: 623-676. edged for loan of material: L. Baert (Museum d’Histoire Bocak L. & Bocakova M. 1990b: Revision of the genus Hemi- naturelle, Brussels), B. Brugge (Zoological Museum, Amster­ conderis (Coleoptera, Lycidae). Acta Entomol. Bohemoslov. dam), T. Huflejt and S. A. Slipinski (Museum and Institute of 87: 209-220. Zoology, Warszawa), M. Kerley and J. Beard (Natural History Bocak L. & Bocakova M. 1991: Revision of the genus Eni­ Museum, London), J. Krikken (Natural History Museum, Lei­ clases C. O. Waterhouse, 1879 (Coleoptera: Lycidae: den), J. J. Menier (Museum national d’Histoire naturelle, Paris), Metriorrhynchinae). Mitt. Munch. Entomol. Ges. 81: K. Matsuda (Takarazuka City, Japan), O. Merkl (Natural His­ 203-226. tory Museum, Budapest), R. Poggi (Museo Civico di Storia Bocak L. & Bocakova M. 1992: Notes on some genera of the Naturale, Genova), W. Schawaller (Staatliches Museum für family Lycidae (Insecta: Coleoptera). Entomol. Basiliensia Naturkunde, Stuttgart), M. Uhlig (Zoologisches Museum, 15: 255-260. Berlin), G. A. Samuelson (State Museum of Natural and Cul­ Bocak L. & Bocakova M. 1999: New taxa of Lycidae from tural History, Honolulu), and H. Schönmann (Naturhistorisches China and Indonesia. Acta Univ. Pal. Olom., Biologica 37: Museum, Wien). 47-56.

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